Ing Nailing and Screwing, and Pre-Boring Is Recommended

UrvlSIANTHE 501 tial of the species in afforestation has been ing nailing and screwing, and pre-boring is recommended. Various quassinoids isolated recommended. The wood is durable, also with- from the plant have shown antimalarial and stands fungal attacks, and is suitable for use in other interesting properties and may have contact with the ground or water pharmacologicalpotential A coumarin derivative has been isolated from Major references Ajaiyeoba & Krebs, 2003; the wood; the bark contains several oxygen- Bolza & Keating, 1972; Burki11, 2000; Dudek, heterocyclic compounds that have not yet been Forster & missenbauer, 1981; Nooteboom, identified. The seeds are rich in mexicanolide 1962; 0teng-Amoako (Editor), 2006; Phong- type Iimonoids and contain 2 triterpenoids. Its phaew, 2003; Saville & Fox, 1967; Stannard, chemical composition confirms the isolated 2000; Voorhoeve, 1979. position of Quiuisiortthe popinoe in the Melt- Other references Adekunle, Oj0 & 01u- aceoe walana, 2002; Adesanwo at a1. , 2004; Ajaiyeoba Botany Deciduous, dioecious, smallto medium- at a1. , 1999; Capuron, 1961; CTFT, 1962e; sized tree up to 20(-30) in tall; hole up to 50 cm Fouarge & G6rard, 1964; Francois at a1. , 1998; in diameter. Leaves alternate, panpinnately Gunb-Fakim & Brendler, 2004; Hawthorne, compound with 5-8 pairs of leaflets; stipules 1995; InsideWood, undated; 1stas, Raekelboom absent; leaflets opposite, elliptical, curieate at & Heremans, 1959; Kryn & Fobes, 1959; Lu- base, obtuse to rounded at apex, entire, pm- monadio at a1. , 1991; Martret, Farmes & Soul. nately veined. Inflorescence a short axillary Ier, 1992; Miralles at a1. , 1988; Neuwinger, panicle, short-hairy. Flowers unisexual, male 2000; Orig, 2001; Perez, 2003; Prot & Korn- and female flowers very similar in appearance, probst, 1985; Takahashi, 1978; Van den Eyn- regular, 5-merous, small; calyx cup-shaped, den, Van Damme & de Wolf, 1994 lobed to the middle; petals free; stamens coin- Sources of illustration Voorhoeve, 1979; PIetely fused into a hairy tube, arithers In- Wilks & Issemb6, 2000. serted along the margin; ovary superior, py- Authors J. N. Gyakari& JR. Cobbinah rainidal, hairy, usuaUy 3-celled, style ending in obscurely 3-lobed stigma; male flowers with rudimentary ovary, female flowers with non- QUrvlSIANTHE PAPiNAE Baill. dehiscing arithers. Fruit a large pyramidal to nearly globose capsule, dehiscing usually with Protologue Grandid. , Hist. phys. Madagascar 3 woody valves, up to 6-seeded. Seeds flat- 34(4), Atlas 2, fasc. 34: t. 251 (1893) tened, with a large wing at apex Family Menaceae Flowering and fruiting are irregular. In west- Origin and geographic distribution Quiui- ern Madagascar ripe fruits are mainly avail- stunthe popinoe is endemic to western and able in November. southern Madagascar. Quiuisionthe comprises a single species. In Uses The wood is used for heavy construc- phylogeny reconstruction based on plastid tion, e. g. for poles of houses and bridges, and DNA sequences, it clustered into the subfamily for heavy carpentry, flooring, cabinet work, and Meltoideoe close to Ekebergio. An unnamed indoor and outdoorJoinery second Quiuisionthe species wlth the vernacu- Properties The heartwood is palebrown, with Iar name 'saniramboanjo' has been reported a purplish tinge when fresh, and distinctly from eastern Madagascar, but it is unclear demarcated from the pinkish, c. 5 cm wide whether it truly belongs in this genus sapwood. The grain Is straight or sometimes The wood of Astrotrichilio, an endemic genus of interlocked, texture coarse. The wood is heavy Madagascar with about 12 species, is often with a density of about 925 kg/ina at 12% inols- known under the same vernacular names as ture content. Shrinkage during drying is high, that of Quiuisionthe popinoe, e. g. 'hornpy'. It is from green to oven dry 5.5% radial and 9.5% lighter in weight, moderately hard and not tangential. Once dry, the wood is somewhat durable, but permeable to preservatives. It is unstable in service. At 12% moisture content, locally used for joinery. However, Astrotrichilio the modulus of rupture is 180 N/mm2, modulus is quite distinct from Quiuisionthe with its of elasticity 21,200 N/min2 and compression stellate hairs and drupe-like fruit parallel to grain 76 N/min2. The wood is very Ecology Quiuisionthe popinoe occurs in dry hard, but not particularly difficult to work with deciduous forest, often along watercourses. It stellite-tipped sawteeth and tungsten-carbide- prefers deeper soils and does not grow well in tipped cutting tools. Splitting is common dur- rocky locations. 502 TIMBERSl

Management Quiuisionthe popinoe usually occurs in scattered groups in the forest. Seed- lings tolerate some shade. However, for proper development and good growth of the seedlings, some thinning or opening of the forest canopy I . is needed. Fruits should be collected from the tree and the seeds sown immediately after col- , Iection for good germination. Genetic resources and breeding Quiui- sinnthepopinoe has been recorded to be locally ,^ common in Madagascar, especially in forests near Morondava. However, little natural vege- L. . tation is left in this region, and Quiuisionthe popinoe may already be subject to considerable genetic erosion. In southern Madagascar it is o also locally common, but grazing livestock and increasing rates of exploitation constitute sen- ous threats. Rhodognupholon breuicuspe - wild Prospects Quiuisionthe popinoe is an jin- portant source of timber for local construction wood, interior joinery, packaging material, in western and southern Madagascar. An in- pallets, boxes, crates, panelling, trim, light ventory of remaining populations of Quiuision- furniture and decorative boards. Traditionally, the popinoe Is needed before it can be deter- canoes are dug out from the hole in Liberia, mined to what extent exploitation of this spe- and smaller branches are hollowed out to pro- cies will remain justified in the future. Virtu- duce quivers in Ghana. The wood is also suit- ally nothing is known about growth rates in able for light construction, light flooring, musi- response to ecological conditions and very little calmstruments, matches, carvings, toys, novel- on natural regeneration. A preliminary study ties, turnery, hardboard and particle board. It indicated that Quiuisionthe popinoe may be is suitable for pulping suitable for enrichment planting. Floss from the fruit is used for stuffing pillows, Major references Gu6neau, Bedel & Thiel, cushions and mattresses, and for making 1970-1975; Randrianas010, 1997; Schatz, 2001 sacks. It has been suggested as a source of pulp Other references Coombes, Mulholland & for paper making. The bark is used for making Randrianarivelojosia, 2005; Muellner at al. , hut walls. A fast, red-brown dye is extracted 2003; Mulholland, Parel & Coombes, 2000; from the bark by boiling and used for dyeing Mulholland & Taylor, 1988; Randrianasolo at cloth. A black dye obtained from the root is a1. , 1996 locally used for making funeral cloth in Ghana Authors R. H. M. J. Lemmens In traditional medicine the powdered root is applied to swellings and dislocations. A paste of the root powder mixed with water is taken to RHODOGNAPHALONBREVICUSPE (Sprague) treat rheumatism and dysentery. The root is Roberty recorded to be used as an antitumour agent in Guinea. A bark decoction is gargled to treat Protologue Bull. Inst. Franc. Mrique Noire, sore throat, and used to promote wound heal- ser. A, 15: 1404 (1953). ing. The bark is also used for treatment of boils Family Bombacaceae (APG: Malvaceae) and venereal diseases, as an emetic, and to Chromosome number2n= 144 prevent abortion Synonyms Bomberx byeuicuspe Sprague (1909), Production and international trade Gabon Bombox cheuolieriPellegr. (1921) exported 3260 ina of Rhodognopholon 67eui- Vernacular names Kondroti(Fr) cuspe timber in 2000, 280 in3 in 2001, 50 ms in Origin and geographic distribution Rhodo- 2002, 790 ina in 2003, 1370 in31n 2004 and 270 gnopholon breuicuspe is distributed from Si. ing in 2005. In 2003 about 3000 in3 was ex- erra Leone eastward to Gabon, Congo and DR ported from Congo, at a price of Us$ 1071m3, Congo, possibly also in Guinea. and in 2005 about 1000 ing at a price of Us$ Uses The wood (trade names: alone, kon- 4031m3. In 2006 Congo exported 16 in3 of logs, droti) is used for the production of veneer, ply- 35 ms of rotary veneer, and 1800 ing of ply-

. RHODOGNAPHALON 503 wood. Cameroon produced 70 ms of this timber in 2000. Properties The heartwood is pinkish to red when freshly cut, turning violaceous brown to brown ochre with darker veins upon drying; it , , is distinctly demarcated from the white and up , . to 10 cm wide sapwood, which turns pale brown on exposure. The grain Is usually straight, texture coarse. 2 The wood is medium-weight, with a density of , .. . .,.. 440-640 kg/ina at 12% moisture content. The rates of shrinkage from green to oven dry are , 4.3-4.7% radial and 4.49.0% tangential. The 5 dry wood is stable in service, but the wood absorbs moisture readily, becoming very heavy, . I , and its use in humid conditions Is not recoin- .,. t, , mended. . ,. , : .. At 12% moisture content, the modulus of rup- ::\ .* ';,* ture is 54-94 N/mm2, modulus of elasticity .I "* , * I " 7100-8800 N/min2, compression parallel to , . , grain 32-45 N/mm2, cleavage 7-16 N/mm, .,. ;! ' *,. 4 Janka side hardness 4540 N, Janka end hard- \ ".. ~ ness 5250 N and Chalais-Meudon side hard- ' ' , $1 - I--' titi ness 1.2-1.8. The wood is moderately brittle ;;' ~~,. .. 3 re#;:;.:&; "..' .,* The wood saws and works easily when sharp- edged tools are used, although it is occaslonally fibrous. It nails and screws well, but holding Rhodognopholon 67euicuspe - I, buse of 601e, ' 2, properties are poor. It does not polish well, but leafy twig, . 3, flowering brunch, . 4, fruit, 5, seed turning and moulding properties are good, and with floss it paints and varnishes satisfactorily. Peeling Redrooon grid odopted by AChmod Satin Nur- characteristics are moderate to good. The wood hornon glues well. It may cause dermatitis in workers when it comes into contact wLth the skin. SIIe, obovate, 3-14 cm x I-4(-5) cm, base The durability is low to moderate; the wood is curieate, apex short-acuminate with obtuse or liable to attacks by fungi, borers and termites notched acumen, margin entire, leathery, dark Both heartwood and sapwood are liable to at- green above, paler green below, glabrescent, tack by Lyetus borers. Both are permeable to pinnately veined with 8-15 pairs of lateral impregnation with preservatives. veins. Inflorescence an axillary 2-3-flowered The wood contains cellulose 43%, pentosans fascicle or flowers solitary. Flowers bisexual, 16%, 11gnin 32% and ash 1.3%. The solubility in regular, 5-merous, fragrant; pedice1 5-10 mm hot water is 2.4%, in alcohol-benzene 2.3% and long, hairy; calyx campanulate, 6-11 mm long, in I% NaOH 18.1% steUate hairy outside, densely to sparsely hairy Description Large, deciduoustree up to 45(- inside, persistent after flowering; petals free, 50) in tall; hole straight, cylindrical, branchless oblanceolate, elliptical or oblong, contorted, for up to 27(-30) in, up to 120(-200) cm in di- 3.5-6 cm x c. I cm, white or pinkish red, hairy ameter, with buttresses up to 1.5 in high and outside, glabrous inside; stamens numerous, c 10-20 cm thick; bark surface grewsh brown, 4 cm long, fused to the petals at their base, smooth on young trees, rough on older trees, filaments grouped in bundles, pink to red; inner bark pink to bright red turning brown on ovary superior, 5-celled, glabrous, style as long exposure, very fibrous, easily detached from as petals. Fruit an obovoid to oblong capsule 5- the wood; crown rounded and lax; buds and 8(-10) cm x 3-4 cm, dehiscent with 5 valves, young branches hairy. Leaves alternate, digi- narrowed at base, apex pointed, glabrous out- tately compound, with 5-7 leaflets, the central side, 4-5-seeded. Seeds pear-shaped, 8-13 mm one largest; stipules linear or triangular, c. 8 x 6-8 mm, glabrous, brown, embedded in min long, densely hairy outside, deciduous; abundant yenowish or reddish brown floss petiole 2-8 cm long, densely hairy; leaflets ses- Seedling with epigeal germination; hypocotyl 504 TIMBERSl

4-7 cm long dispersed by wind together with the floss. Other botanical information Rhodogno- Rhodognopholon breuicuspe is characterized as photon comprises 3 species, all in tropical at- a light-demanding pioneer, but regeneration is rica. It was formerly included in Bombox, but it rarely abundant. is now considered a separate genus, differing Ecology Rhodognopholon breuicuspe occurs from Bombox in being unarmed, and having scattered in primary as well as secondary for- larger seeds, only I whorl of stamens and per- est, but Is most common in secondary ever- sistent calyx green forest Anatomy Wood-anatomical description OAWA Propagation and planting The 1000-seed hardwood codes): weight is about 100 g. Germination normally Growth rings: 2: growth ring boundaries iridis- starts after 5-8 days, with 60-80% germina- tinct or absent. Vessels: 5: wood diffuse-porous; tion. It is recommended that nursery-grown 13: simple perforation plates; 22: intervessel seedlings not be planted out until they are at pits alternate; 23: shape of alternate pits po- least I year old. Iygona1; 26: intervessel pits medium (7-10 pin); Management In south-western Garnero0n 27: intervessel pits large G 10 prn); 31: vessel- Rhodognopholon breutcuspe trees with a di- ray pits with much reduced borders to appar- ameter over 60 cm were found at a mean den- ently simple: pits rounded or angular; 43: mean sity of only 0.02 trees/ha, with an average tangential diameter of vessellumina z 200 grn; standing hole volume of 0.14 ms per ha. In 46: S 5 vessels per square minimetre; (47: 5-20 western Gabon the estimated standing volume vessels per square minimetre); 56: tyloses in the 1960s was 0.17 ina per ha. common. Tracheids and fibres: 61: fibres with Diseases and pests Rhodognopholon breui- simple to minutely bordered pits; 66 non- cuspe is a host tree of Cacao Swollen Shoot septate fibres present; 69: fibres thin- to thick- Virus (CSSV) which causes swollen shoot dis- walled. Axial parenchyma: 77: axial paren- ease in cocoa, a disease that has had a devas- chyma diffuse-in-aggregates; 79: axial paren- tating effect on cocoa production in Ghana and chyma vasicentric; 86: axial parenchyma in neighbouring countries. The virus causes chio- narrow bands or lines up to three cells wide; rosis of the leaves in Rhodognopholon breui- 92: four (3-4) cells per parenchyma strand; 93: cuspe. eight (5-8) cells per parenchyma strand; (94: Harvesting In C6te d'TVoire the minimum over eight cells per parenchyma strand). Rays: cutting diameter is 60 cm, in Ghana 70 cm. 97: ray width I-3 cells; (102: ray height > I Yield It has been estimated that trees with a mm); 104: all ray cells procumbent; 106: body hole diameter of 60 cm, 70 cm and 100 cm yield ray cells procumbent with one row of upright 4.3 ina, 5.8 ina and 11.9 ms of wood, respec- and/or square marginal cells; 1/5: 4-12 rays tively. per mm. Stoned structure: (119: low rays sto- Handling after harvest After felling, rapid ried, high rays non-stoned); 120: axial paren- extraction from the forest or treatment with chyma and/or vessel elements stoned. Mineral preservatives Is necessary to prevent deter10ra- inclusions: 136: prismatic crystals present; 141 tion of the wood by fungiand insects. prismatic crystals in non-chambered axial pa- Genetic resources Rhodognopholon 67eui- renchyma cells. cuspe has a wide distribution, but is nowhere (P. Mugabi, A. A. Oteng-Amoako & P. Baas) abundant. It is classified as vulnerable in the Growth and development Initial growth of JUGN Red list, as its occurrence is decreasing Rhodognopholon breuicuspe is very slow. The and it is overexploited mean height of nursery-grown plants in Prospects The wood of Rhodognopholon bre- Guinea was 13 cm at 3.5 months after sowing, uicuspe has low strength and is not durable, 20 cm after 10 months and 80 cm after 2 years but the species has local importance as a inul- After planting out in the field, the growth was tipurpose tree and commercial importance as a 20 cm/year during the first 2 years. In southern source of plywood. Given its slow growth it is C6te d'TVoire the average annual diameter in- unlikely to gain importance as a plantation crement was 2.4 min in natural evergreen for- species. In view of its vulnerable conservation est and 4.3 min in thinned forest. In Liberia, status, natural stands should only be allowed C6te d'Ivoire, Ghana and Cameroon flowering to be exploited in a sustainable way is usually in November, and fruiting in Febru- Major references Beentje & Smith, 2001; ary-March. Flowers develop in the dry season, Bolza & Keating, 1972; Burki11, 1985; CTRAD often when the tree is leafless. The seeds are Forestry Department, 2003; G6rard at al. ,

. RHODOGNAPHALON 505

1998; Takahashi, 1978; Thiraku1, 1983; when freshly cut, but on drying the approxi- Viniers, 1973b; Vivien & Eaure, 1985; Voorhoe- mately 7.5 cm wide sapwood becomes cream- ve, 1979. coloured. The grain is straight, texture medium Other references Adu-Gyamfi, 2006; to moderately coarse. Black gum ducts are ATIBT, 1986; ATIBT, 2005; ATIBT, 2007; sometimes present. Beentje, 1989; CTFT, 1961e; de in Mensbruge, The density of the wood is (420-)465-480(-580) 1966; de Saint-Aubin, 1963; Durrieu de Mad- kg/ing at 12% moisture content. The wood dries ron at a1. , 1998a; Gassita at al. (Editors), 1982; rapidly with some surface checking and distor- Graham at a1. , 2000; Hawthorne, 1995; Haw- tion, and occasional collapse. The rates of thorne, 1998c; ICTV, undated; InsideWood, shrinkage from green to 12% moisture content undated; ITT0, 2006; Kryn & Fobes, 1959; are 2.0% radial and 3.5% tangential. Star Neuwinger, 2000; Raponda-Walker & Sillans, shakes develop on drying. Once dry, the wood 1961; Viniers, 1975b; Wilks & TSSemb6, 2000 is fairly stable in service Sources of illustration Viniers, 1973b; The wood is weak and soft. At 12% moisture Voorhoeve, 1979; Wilks & TSSemb6, 2000 content, the modulus of rupture is 47 N/min2, Authors V. A. Kerneuz6 modulus of elasticity 6100 N/mm2, compression parallel to grain 31 N/mm2, shear 6 N/mm2, cleavage 44 N/min radial and 53 N/min tangen- RHODOGNAPllALONSCHUMANNIANUM tial, and Janka side hardness 2000 N A. Robyns Both the green and dry wood saw easily. The wood works easily with hand and machine Protologue Bull. Jard. Bot. Etat 33: 263 tools. It takes nails well, but the holding power (1963) is low. It peels easily. Painting, staining and Family Bombacaceae (APG: Malvaceae) lacquering properties are poor. Veneers dry SynonymsBomboxrhodognupholonK. Schum rapidly, with high shrinkage, and tend to be (1895) brittle. Vernacular names East African bombax, The wood has low durability, being liable to wild kapok tree (En). Msufi inwitu, msufi pori attacks by sapstain fungi, marine borers and (Sw) termites. The sapwood is liable to attack by Origin and geographic distribution Rhodo- Lyctus borers. The heartwood is extremely re- gnupho!on schumonnionum is distributed in SIStant to impregnation with preservatives Kenya, Tanzania, Malawi and Mozambique. It The wood fibres have an average length of 1.9 is occasionally planted mm, with a diameter of 25.4 prn. The chemical Uses The wood is locally used for roofing, composition is: cellulose 50%, pentosans 8%, doors, panelling and cheap plywood for packing 11gnin 35% and ash I%. The solubility in cold cases. The hole is traditionally used for dugout water is I%, in hot water 2%, in alcohol- canoes. The wood is suitable for low-grade fur- benzene 4% and in I% NaOH 16%. Paper- niture, sporting goods, matches, hardboard, making experiments in the 1950s resulted in particle board and wood-wool. It is also suit- papers with low strength able for pulping and for charcoal making Ethanol, petroleum ether and ethyl acetate The roasted seeds are eaten like groundnuts, extracts of the root bark have shown in-vitro and they are pounded and cooked with vegeta- antimalarial activity bles or meat. Floss from the fruit has been used Botany Medium-sized to fairly large tree up for stuffing pillows, cushions and mattresses to 40 in tall; hole cylindrical, branchless for up The bark yields a red-brown dye and fibre for to 21 in, up to 150 cm in diameter, buttresses rope making. The roots are used to treat up to 3 in high; bark surface smooth, yellow- asthma, coughs and diarrhoea, the bark is a green, on older trees sometimes scaly and grey traditional medicine against diarrhoea and Leaves alternate, digitately compound, wlth malaria, and the leaves are used in cleansing (3-)5-7(-8) leaflets, central leaflet larger than rituals. Rhodognopholon schumonnio, ,urn is other ones; stipules deciduous; petiole 3.5-12.5 used as a shade and wayside tree cm long; petiolules 0.5-2 cm long; leaflets ellip- Production and international trade The tical or obovate, 3-14 cm x 2-6 cm, curieate or seeds are sold in local markets decurrent at base, acuminate at apex, margin Properties The heartwood is pale to dark entire, glabrous or steUate hairy, pinnately pinkish brown with diffuse dark bands; it is veined with 7-22 pairs of lateral veins. Inno- indistinctly demarcated from the sapwood rescence an axillary, 2-5-flowered fascicle, or 506 TIMBERSl

flowers solitary. Flowers bisexual, regular, 5- insects and fungi. Seedlings are planted out merous; pedice1 6-25 mm long, glabrous or when they are about I year old and 60-120 cm stellate hairy; calyx campanulate, truncate or tall. Wild trees are locally protected. In Mo- slightly lobed, 6-14 mm x 7-11 mm, glabrous zambique the minimum felling diameter is 50 or stellate hairy outside, silky hairy inside; cm, In Tanzania 55 cm petals 5-11 cm x I-6 cm, obtuse to acute at Genetic resources and breeding It is un- apex, red, pale yellow or white, hairy on both clear to what extent Rhodognopholon schu- sides; stamens numerous, 4.5-7 cm long, fused monnionum is threatened with genetic erosion. to the petals at their base, united at their base It has been recorded as being at low risk I of Into 5 bundles, red; ovary superior, ovoid, 2-4 least concern, and it Is not included in the min long, hairy, 5-celled, style 5-11 cm long IUCN Red list Fruit an ellipsoid to obovoid capsule 5-13 cm x Prospects Rhodognopholon schumunnionum 2.5-4.5 cm, dehiscent with 5 valves, hairy or is a multipurpose tree, yielding not only wood, glabrescent, brown, many-seeded. Seeds glo- but also food, fibre and traditional medicines. hose or angular, 8-11 min in diameter, gla- The wood has poor strength and durability brous, brown, embedded in reddish brown floss. characteristics, however, and the importance of Seedling with epigealgermination. Rhodognopholon schumonnionum as a source Rhodognopholon comprises 3 species, allin of timber Is unlikely to increase tropical Africa. It was formerly included in Major references Beentje & Smith, 2001; Bomberx, but it Is now considered a separate Bolza & Keating, 1972; Bryce, 1967; Ruffo, genus, differing from Bomberx in being un- Birnie & Tengnas, 2002; Takahashi, 1978 armed, and having larger seeds, only I whorlof Other references Beentje, 1989; Beentje, stamens and persistent calyx. 1994; de Seabra & Perreirinha, 1950; Dowsett- Within Rhodognopho!on schumonnionum 2 Lemaire & White, 1990; Gessler at a1. , 1994; varieties are distinguished: Lovett at a1. , 2006; Lutze, 2001; Pakia & - var schumonnionum (synonyms: Bombox Cooke, 2003b; Parry, 1956; Williamson, 1955 rhodognopholon K. Schum var rhodogno- Authors M. Brink photon, Rhodog, ,opholon tongonyikense A. Robyns), with leaves and pedicels glabrous; distributed in Kenya, Tanzania and SanOANAIAMADAGASCARIENSIS R. Vig. Mozambique; - var. tomentosum A. Robyns (synonyms: Born- Protologue Notul. Syst. (Paris) 14(3): 186 box rhodognopholon K. Schum. var. tomento- (1951) sum A. Robyns; Bombox mossombicense Family Papilionaceae (Leguminosae - Papil- A. Robyns; Bombox stolzii Ulbr. , Rhodogno- ionoideae, Fabaceae) pholon mossombicense (A. Robyns) A. Robyns; Origin and geographic distribution Sohoo- Rhodognopholon 8101zii colbr. ) A. Robyns), riglo mudogoscoriensis is endemic to eastern with leaves and pedicels sparsely to densely Madagascar, where it is restricted to the stellate hairy; distributed in Mozambique. coastal zone around Toamasina Initial growth of Rhodognupholon schumon- Uses The wood is highly valued for house itionum is fairly fast. Three years after sowing, construction seedlings may be 2-3 in tall. In Tanzania the Properties The wood is yenowish, with a fine fruits ripen in October-December. The seeds texture. It can be polished well are dispersed by wind, but most seeds do not Botany Deciduous small to medium-sized tree spread further than 100 in from the tree up to 15 in tall; hole up to 50 cm in diameter; Ecology Rhodog, lopholon schumonnionum bark bronzed brown, smooth, with greyish sur- occurs in wooded grassland, woodland and for- face cracks; twigs thick and succulent, shortly est, from sea-level up to 1100 in altitude. The hairy when young. Leaves alternate, imparip- average annual rainfall in the area of distribu- innate with 9-13 leaflets; petiole and rachis tion is normally notlessthan 1000 mm. channelled, sparsely shortly hairy; leaflets Management Rhodognopho!on schumon- opposite to slightly alternate, oblong-elliptical itionum is easily propagated using seed. The to ovate, (2.5-)3-7.5 cm x (I-)1.5-3.5 cm, 1000-seed weight is about 270 g. Fresh seeds rounded to truncate at base, obtuse to shortly normally germinate well, with germination acuminate at apex, thlnly leathery, glabrous taking 4-21 days. The seeds can be stored for except for the inIdrib and margins. Innores- up to 4 months, but are liable to attacks by cence a rather lax raceme 5-23 cm long, many-

. SIDEROXYLON 507 flowered. Flowers bisexual, papilionaceous; used for house construction and for building pedicelwith smallbract near the middle; calyx boats, bridges and mills. It is also used as fire- broadly cup-shaped, 5-7 mm long, with wood and for charcoal making rounded lobes; corolla violet, with circular The fruits are sometimes eaten. In traditional standard c. 2 cm in diameter, long-clawed at African medicine the roasted powdered root is base, notched at apex, wings and keel oblong, mixed with oil from the seed of TrLchilio shorter than standard; stamens 10 or 11, free; emetico Vahl and rubbed into incisions over ovary superior, shortly stalked, glabrous, I- fractured limbs. A decoction of the root, admin- celled, style slightly UPCurved, stigma Iridis- 1stered as an enema, is a diaphoretic. The dried tinct. F"uit an oblong pod (4-)6-14 cm x 1.5-3 pulverized root is eaten to treat conjunctivitis. cm, flattened, with slightly wing-like margins, Minfusion of the bark is taken against night- glabrous, indehiscent, (I-)3-5-seeded. Seeds mares. A decoction of the bark is given to treat kidney-shaped, c. 7 min long, flattened, chest- gall sickness in animals. nut-brown Properties The wood is yenowish brown with Soh00,1010 comprises 2 species, both restricted a fine texture. It is heavy (density 1040 kg/in3 to Madagascar. The genus is related to Neo- at 10% moisture content), hard, strong and hormsio, which differs in the usually smaller durable, even in damp circumstances number of leaflets per leaf, campanulate calyx Botany Spreading, much-branched, evergreen and dehiscent pods shrub or smalltree up to 15 in tan, often with a Sohoonolo mudugoscoriensis is tardily decidu- gnarled appearance, with scarce milky latex; ous. It usually flowers on leafless shoots Just bark grey, brown or black, fissured; young after the leaves have fallen. It is reported to fix branches covered with fine soft grey to rusty atmospheric nitrogen hairs, older branches glabrous. Leaves spiralIy Ecology Sohoonolo mudogoscoriensis occurs arranged or less frequently opposite, simple in humid coastal forest on sandy soils. and entire; stipules absent; petiole 05-2 cm Genetic resources and breeding Sohoo- long, rusty hairy when young, glabrous later; ito10 madogoscoriensis is classified as endan- blade elliptical to obovate, (3-)4-12(-15) cm x gered in the 2006 IUCN red list of threatened (1.5-)2-6(-7.5) cm, base narrowly curieate, species. It is selectively felled for timber and apex obtuse to rounded or notched, thickly the remaining fragments of coastal forest in its leathery, often with rusty hairs disappearing small distribution area are still declining. with age, pinnately veined with indistinct Iat- Prospects Protection of this threatened spe- eral veins. Inflorescence a congested fascicle in cies is badly needed and harvesting should be leaf axils. Flowers usually bisexual, regular, 5- stopped immediately to preventits extinction merous, with an unpleasant smell; pedicel up Major references du Puy at a1. , 2002 to 7 min long, shortly hairy; sepals broadly Other references du Puy & Labat, 1998k; ovate, up to 2.5 mm long, slightly hairy or gla- Peltier, 1972; Schatz, 2001 brous outside; corolla campanulate, up to 5 min Authors R. H. M. J. Lemmens in diameter, whitish, cream-coloured or greenish, tube up to 1.5 mm long, lobes ovate and up to 2.5 min long; stamens opposite corolla lobes, SIDEROXYLONINERME L. up to 5 mm long, alternating with petalold, ovate to lanceolate staininodes 1.5-3 min long; Protologue Sp. PI. I: 192 (1753). ovary superior, globose, hairy, 5-eeried, style up Family Sapotaceae to 1.5 mm long. Fruit a globose berry 6-15 mm Chromosome number2n=44 in diameter, with persistent style, ripening Vernacular names White milkwood (En). purplish black, smooth, pulp fleshy, sticky, I- Mkoko bara, ingongonga, intunda wa rigombe, seeded. Seed globose, 5-9 min in diameter, inchocha inwitu (Sw). testa thick and woody, cream-coloured or Origin and geographic distribution Sidero- brown, shiny, with 5 longitudinal ridges and 2- xylon titerme is distributed along the eastern 4 smallpits near the basal scar coast of Africa, from Somalia southwards to In southern Africa Sideroxy!on titerme flowers South Africa, and on kidabra Island (Sey- in JanuaryJuly; fruiting is in JulyJanuary. chenes) and Mayotte; it occurs sporadically Seed dispersal is by birds. Sideroxylon titerme more inland. can become very old: the tree known as 'Post Uses The wood of Sideroxylon titerme is used Office tree' in Mossel Bay (South Africa) is for poles and for making spoons, and has been more than 500 years old.

. 508 TIMBERSl

Within Skierory!on titerme 3 subspecies are Other references Beentje, 1994; BOSman, distinguished: subsp. diospyroides (Baker) 2006; Friedmann, 1981; Hemsley, 1968; J. H. Hemsl. , distributed in Somalia, Kenya, Holmes & Cowling, 1993; Neuwinger, 2000; Tanzania, Zimbabwe and Mozambique; subsp Sharma & Singh, 2002; Turpie, 2000; van titerme, occurring in Mozambique and South Vuuren, Banks & Stohr, 1978; van Wyk & van Africa; and subsp. cryptophlebium (Baker) Wyk, 1997 J. H. Hemsl. occurring on A1dabra Island (Sey- Authors M. Brink chelles) Sideroxylon comprises about 50 species in tropical America and about 25 species in the SINARUNDINARIAALPINA (K. Schum. ) Old World (6 in mainland Africa, about 6 in CS. Chao & Renvoize Madagascar, 8 in the Mascarene islands, and 5 in Asia). Skierory!Dripuberulum DC. , a tree up Protologue Kew Bull. 44(2): 361 (1989) to 15 in tall with a hole diameter up to 60 cm, Family Poaceae (Gramineae) is endemic to Mauritius, where it is known as Synonyms Arundinori0 o4pino K. Schum. 'mangher rouge'. Its wood is heavy and durable (1895), Yushonio ulpino (K. Schum. ) W. C. Lin and is used for making poles, planks and keels (1974). of boats. The wood of Sideroa:y!on sessili/10rum Vernacular names African alpine bamboo, (Poir. ) Capuron, a rather rare endemic ofMau- mountain bamboo (En). Bambou creux (Fr). ritius, has been described as very good for Mianzi, inwarizi(Sw) cabinet work Origin and geographic distribution African Ecology Sideroxylon titerme is essentially a alpine bamboo occurs in scattered populations tree of coastal woodland and littoral forest. It is on mountains from southern Sudan and Ethio- a common component of shrub thickets on the pia southwards to Malawi. hamterval of 2000 seashore near the high-water mark and is kin separates its occurrence in western Camer- found along landward fringes of mangroves. It o0n from those in eastern Africa. It is fre- sometimes occurs further inland along rivers quently planted, e. g. in Ethiopia and Zim- and in open woodland, up to 1500 in altitude, babwe. often on termite mounds. It occurs in regions Uses The whole stemsofAfrican alpine bam- with an average annual rainfall of 300-1500 boo are used for hut construction, particularly mm and tolerates shade andwind. as rafters, and for fencing. In the Foroto moun- Management Skierorylon merine is easily tains in Tanzania entire villages are con- propagated by seeds, which take 4-6 weeks to structed using this bamboo, and water pipes germinate. Vegetative propagation using cut- made from African alpine bamboo supplied tings is also possible, but only semi-mature water to around 100,000 people in the 1980s. A side shoots should be used; cuttings normally few specialized enterprises produce bamboo rootin 6-8 weeks. furniture. Split canes are woven into storage Genetic resources and breeding It is un- pots and baskets in Tanzania and Kenya; in clear to what extent Sideroxylon merine is threatened by genetic erosion in tropical at- rica. It Is protected in South Africa, where even for pruning a permit is required; 3 specimens have been declared National Monuments in . South Africa. Prospects The wood of Skierorylon merine is hard, strong and durable, but little information on the wood properties is available. In view of , the small size of the tree and the uncertain .^ conservation status of the species, increased Importance as a source of tlruber is not to be .. expected. Skierory!on titerme does not easily catch fire and plantings could be useful as a firebreak o Major references Coates Palgrave, 1983; Friis, 2006; Kupicha, 1983; Lovett at a1. , 2006; Pennington, 1991 Sinorundinorio ulpino - wild

. SINN{UNDINARIA 509

Uganda they are made into beehives. Dry stems are used as fuel. Young shoots are mar- keted as a vegetable 000unt Elgon, Kenya/Uganda I I^ border), and leaves and fine branchlets serve as cattle fodder. Sinorundinori0 o10ino is also .I . valued in soil conservation as a more effective \ watershed coverthan trees \. 1.1^I '** Production and international tradeAfrican *.. I .I alpine bamboo forest is most extensive in the 3 I high mountains of the Nile/Congo divide and ,. further east (500,000 ha, with over 100,000 ha in each of DR Congo, Ethiopia and Kenya). In .t \ Cameroon the area is less than 20,000 ha. Its gregarious character typically results in a standing crop of more than 5000 full-sized I stems per ha in mixtures with Afromontane 11. trees, and up to 40,000 stems per ha in pure I stands. There is no international trade based on the species, butsome commerce exists in the countries where it grows. Levels of utilization reported in national statistics, based on pay- merits for harvesting licences, underestimate 2 exploitation for immediate local community needs. Properties The mean density of the stem Sinorundinorio ulpino - I, led/y brunches, . 2, wall of African alpine bamboo is about 0.7 flowering brunches, . 3, spikelet, . 4, groins g/cm3 at 8% moisture content. Dried stems Redrown und adopted by Achingd Satin Nur- used in construction and fencing are susceptl- hornon ble to infestatton by the powder-post beetle Dinoderus minutus. Nevertheless, the stems to 20 in tall, up to 12.5 cm in diameter, yellow- are considered durable and houses and fences ish at maturity, hollow, thick-walled, many- made from them in DR Congo are said to last noded, profusely branched. Leaves alternate, for more than 20 years' simple; stern leaves with sheath ovate to ob- Stems from DR Congo contained: bolocellulose long-lanceo}ate, glabrous or with reddish 60-65%, u-cellulose 49%, pentosans 17%, 11g- brown bristly hairs, tipped with a linear blade rim 24%, ash 3%. Solubilities were 3.6% Clot c. 6 cm long; branch leaves with sheath having water), 1.8% (alcohol-benzene) and 22.2% (I% smalllateral auricles, 11gule c. 2 mm long, NaOH). The mean fibre length was 2.0 mm, blade linear-lanceolate to narrowly lanceolate, with a diameter of 17.9 prn, a lumen width of 5-20 cm x 0.5-1.5 cm, base constricted into a 3.6 prn and a cell wall thickness of 7.2 pin short stalk, apex acute or with apicalbristle up Sterns give reasonable yields of readily to 2 cm long, glaucous to bright green, gla- bleached pulp suitable for producing writing brous, with conspicuous transverse veins. In- and printing paper. norescence a terminal panicle 5-15 cm long. The foliage is fibrous (crude fibre 26.6%) and Spikelets linear to linear-lanceolate, 15-50 mm rich in ash (15.3%). analysis of material from x 3-4 mm, comprising 4-11 florets, the apneal Ethiopia onasha) indicates that stems are low ones sterile; glumes lanceolate or narrowly in N, particularly those more than 3 years old ovate, 4-8 mm long, apex acuminate, papery, co. 3%), but in leaves/branchlets the level is 5-9-veined, persistent; lemma lanceolate- higher (1.8%). Rhizomes are relatively rich in oblong to ovate, 7-10 min long, apex acute, 7- potassium (1.4%) and leaves/branchlets much 9-veined, palea linear-oblong, about as long as richer in calcium (0.3%) than other parts of the lemma, apex truncate, 7-9-veined; norets with plant (0.02-0.06%). PhDsporus levels are fairly 3 stamens, 3 10dicules, and glabrous ovary with uniform throughoutthe plant (0.05-0.16%). 2 stigmas. Fruit a spindle-shaped caryopsis Description Evergreen bamboo with short (grain) 1.5-6 min long, blackish brown, with a rhizome up to 10 cm thick, and usually not longitudinal furrow. closely clustered stems; stem (CUIm) erect, up Other botanical information Sinoru, Idi-

. 510 TIMBERSl norm comprises about 50 species, most of them Ecology Sinorundinori0 o10ino is restricted from tropical Asia, 2 in Central America, I in to high elevations (2000-4000 in altitude) and mainland Africa and 2 in Madagascar. Sino- is the characteristic and definitive dominant of rundinori0 o40ino has been included in several Afromontane bamboo vegetation. It also occurs genera, but in recent molecular phylogenetic in abandoned fields and it can form extensive analysis its exact position within the so-called pure stands. Afromontane bamboo vegetation Thornnocolomus group and allies remained occurs in cool growing conditions, with average unclear. Additional research is needed. annual temperatures of 14-17'C. Average In the past, smallindividuals of African alpine monthly maximum temperatures are 13-32'C, bamboo have occasionally been misidentified as and average monthly inlnimum temperatures Thornnocolomus tesse!lotus (Nees) Soderstr. & range from -4'C to 11'C, implying that some R. P. Ellis, the South African mountain bamboo. populations tolerate frost. Rainfallis seasonal, Thornnocolomus tessel!otus differs in having with 3-6 dry months (mean rainfallless than thin-walled sterns arising from elongated rhi- 50 mm) in eastern Africa, but only 2 dry zomes, stem leaves with lanceolate blades and months in Garnero0n. annual totals vary from without auricles, and branch leaves with hard 800 min in Tanzania to 2000 min in Ethiopia acuminate tips and hairy 11gule, whereas its and 3000 min in Cameroon. Climate require- spikelets are sessile and ovate, and contain merits over-ride soiltype requirements, with only a single fertile floret. occurrences on impoverished ferralsols, moder- Growth and development Although nurs- ately fertile cambisols, and richer andosols and ery seedlings have been raised in Kenya, there nitisols. Well-drained humus-rich soil on gentle is no published description of the germination slopes and in ravines, with space for vigorous and initial growth of African alpine bamboo rhlzome development, allows luxuriant growth. Small plants, consisting entirely of new shoots, On shallow soils and rocky ground individuals form loose, but discrete, clumps. With time, are stunted. Afromontane bamboo forest has sterns at the centre die and disappear, while been descrlbed as a climax formation, but new stems are added at the periphery. Re- burnt forest tree stumps within Sinorundi- peated exposure to fire causes mortality of nori0 o1pino stands have been interpreted as young peripheral rhizomes and concentration evidence of a fire-induced community. Another of new shoots close to older ones, maintaining view treats the species as a light-demanding relatively compact clumps. Rhizome networks pioneer forming populations maintained by producing new stems annually may survive for large herbivore activity. at least 40 years, but individual stems survive Propagation and planting Attempts to only 8-14 years. New stem production is a sea- germinate seed of African alpine bamboo from sonal process, allowing cohorts of stems in dif- the few seed crops occurring are not always forent age classes to be recognized. A year of successful, but in Kenya seeds sown in nursery vigorous stem production may be followed by a beds and watered daily have germinated. Seed- 2-3-year period of low stem production. Un- lings 2-3 cm tall were transferred to soil boxes usual droughts also reduce production in the and planted out 8-12 months later, at 2 in subsequent growing season The annual spacing. A stand with stems up to 12 in tall growth cycle involves the development of new and 5 cm in diameter resulted after 6 years stems at rhizome apices where growth is acti- In parts of Ethiopia and Uganda offsets are vated when the rainy season starts. New stems often planted. In experiments in Kenya, there reach full height in 2-4 months and branch in has been successful use of offsets (single stems the following year. By the fourth year of shortened to 60 cm, with attached rhizome), growth the stems are glabrous and sufficiently clump division (groups of 5 stems shortened to rigid for use as poles. Estimates of the interval 60 cm, with the parent rhizome) and 20 cm between flowering events vary from 15 years lengths of rhizome. Offsets with stems pro- (Mount Elgon, Kenya) to 40 years (Aberdares duced in the previous growing season are pre- Range, Kenya). Flowering may be synchronous forred. Stern cuttings have not produced in patches several hectares in extent within a shoots, not even after treatment with rooting population. It is generally assumed that plants compounds. die after flowering, although development of Management Management of African alpine new shoots from parts of the rhizome network bamboo is mainly limited to harvesting of surviving after flowering has been noted in natural stands, undertaken by area rather Kenya. than by clump, because stems are usually well

. SINN{UNDINARIA 511 separated. For Kenya, burning after exploita- population of Chimaliro, Malawi, is thought to tion has been recommended. For nitisols in the be recently extinct. Other populations have Masha Forest in Ethiopia, applications of P shrunk, with causes suggested being succes- and K at the end of the rainy season have been SIonal vegetation change following fire, exclu- recommended to sustain productivity SIon and elimination of elephants and buffaloes Diseases and pests Association of African (EChuya, Uganda), conversion to agricultural alpine bamboo with the hasidiomycete Armil- land (Ethiopia) and over-exploitation of the loner meneo has been reported in Kenya and sterns (Tanzania, Uganda). there is suspicion that the fungus spreads from Prospects African alpine bamboo is of high a reservoir in the bamboo to planted pines and local importance throughout its range, and in hardwood trees Ethiopia, Kenya, Tanzania and Uganda there Harvesting Natural stands of African al- is a consistent smallsupply of bamboo material pine bamboo can be clear-felled but recovery is and products to markets remote from the slow, development offun-sized stems taking 9- source areas. The International Network for 10 years. Stems must be full-sized and at least Bamboo and Rattan (INBAR) has evaluated 3 years old before they can be exploited for bamboo 'production to consumption systems in structural use, and numbers must accumulate these 4 countries. Exploitation is not subject to to levels making harvesting worthwhile, so effective regulation and monitoring, and de- felling cycles of 14-21 years have been recoin- clared policies on national bamboo resources mended. The cycle can be reduced to 5-6 years are lacking. There is little professional exper- on good sites if modest yields are acceptable tise to promote active management and refine and 50% of the mature stems are retained harvesting to obtain shoots as well as young Harvesting of stems under 2 years old suitable and mature poles, or encourage the develop- for weaving, and harvesting of edible shoots as merit of processing and marketing frameworks vegetable are rainy season activities. Nevertheless, enough SIIvicultural knowledge Yield The dry weight of a standing crop of exists for preliminary management schedules stems of well-stocked stands in Masha, Ethio- to be applied, taking account of the age struc- pia has been estimated at 51tftia and in Kenya ture of the stands in the context of marketable at 97 tma. With a 5-year cutting cycle exploit- products and harvesting intensity and fre- ing only mature stems, representing 20% of the quency. The infrequent flowering and uricer- number of stems present, a potential yield of tain seed viahinty invite development of an 10 tma per year has been estimated for Ethio- economicalIy attractive method of vegetative pia. Better management might raise this to 15 propagation, for stand rehabilitation and wider t/ha per year planting on agricultural land. The distinctive Handling after harvest Stems for construc- dominance, or even purity, of the species where tion are stripped of branches and trimmed to it grows, and the short-rotation potentiallus- lengths of 7.5-9 in. Drying and protection tify further research into prospects for bamboo against the powder-post beetle Dinoderus material entering the pulp/paper industry if minutus attack are advisable. Protection of demand exists and if sufficient material of complete stems with preservative solutions Is quality coinparable or superior to current al- difficult but it is assumed that soaking in wa- ternatives can be sustainably supplied. Clanfi- ter for 2-3 months, widely used with other cation of geographic variation through co- bamboos, may provide some protection ordinated systematic study could reveal prove- In Uganda internodes of stems harvested for nances of differing quality and relevance for weaving material are cut into slivers which Improvement Inltiatives may be bundled and stored for several months Major references A1vin0, 1950; Clayton, before use. Edible shoots are sun-dried or 1970; Clayton, Harman & Williamson, 2002- a; smoked and can be stored for up to 2 years' Binbaye at a1. , 2005; Hemp, 2006; Hubbard, Genetic resources Today's distribution of 1962; 1stas & Raekelboom, 1962; ingomo & Sinoruiidinorio dipino, including intervals Kamiri, 1987; Launert, 1971; Wimbush, 1945 well in excess of 100 kin, makes geographic Other references Ayre-Smith, 1963; Ba- differences probable. There are no germplasm nana & Twehey0, 2001; Banana & Tweheyo, collections, however, although ex-situ cultiva- 2004; Chao & Renvoize, 1989; CTFT, 1962a; tion is reported for the Muguga Arboretum, Cunningham at a1. , 1970; BSegu, Ssenteza & Kenya. Reports at national levelindicate popu- Sekatuba, 2000; Gibson, 1960; Guo & Li, 2004; Iation losses and conservation threats. The Kelbessa at a1. , 2000; Kigom0, 1990b; Li, 1997;

. 512 TIMBERSl

Lovett at a1. , 2006; Mgeni & Swai, 1982; Ohm- yellow-brown stellate hairy; bracts c. 7 mm x 3 herger, 1999; Origugo at a1. , 2000; Phillips, mm, caducous. Flowers unisexual, regular, 5- 1995; Snowden, 1953; Soderstrom & Ellis, merous, yellowish, c. 7 mm long; perianth cain- 1982; van der Zori, 1992. panulate with lobes about as long as tube, stel- Sources of illustration Clayton, 1970; late hairy; male flowers with 8-10 arithers Engler, 1908 borne on a long common stalk; female flowers Authors J. B. Hall& T. Inada with ovary consisting of 5 carpels united loosely. Fruit consisting of 4-5 woody follicles (5-)7-9 cm x 3.5-6 cm, rusty brown-stellate STERCULIAAPPENDICULATAK. Schum. hairy, many-seeded. Seeds ovoid, 1.5-2 cm x I- 1.5 cm, glabrous, pale yellow, dangling on a Protologue Engl. , Pftanzenw. OSt-^ticas O white thread I-2 cm long. 271 (1895). Sterculto comprises about 150 species and oc- Family Sterculiaceae (APG: Malvaceae) CUTS throughout the tropics. In tropical Africa Vernacular names Tall sterculia (En) about 25 species can be found. 8/@reulio up- Mfune, ingude, msefu (Sw) pendiculoto is often confused with Sterculio Origin and geographic distribution Stercu- qttinquelobo (Garcke) K. Schum. The latter nor OPPendtculoto is distributed from Kenya species has a flaking bark, occurs in drier re- south to Malawi and Mozambique. In Zim- gions, and is found at altitudes up to 1650 in. babwe it is rare and restricted to the extreme The differences in flowers and fruits between north of the country. the 2 species are distinct Uses The wood of Sterculio OPPendiculoto is Sterculio OPPendiculoto grows fairly fast. In used for local construction, boxes and plywood Tanzania seeds ripen in August-September. In It is also used as firewood large parts of its range in Tanzania Sterculio Cooked leaves are eaten as a vegetable. Seeds OPPendicu!oto is said to be valued for tradi- are collected from the ground, roasted and tional ceremonies or is considered sacred and eaten whole or pounded, and cooked with vege- usually left standing like baobab (Adonsonio tables. Unspecified plant parts yield a yellow- digitoto L. ) trees when clearing land for culti- brown dye that is used by the Shambaa people vation of Tanzania. A decoction of the roots is drunk Ecology Sterculio OPPendiculoto is found in to prevent miscarriage, to cure diarrhoea and coastal and lowland riverme forests from sea- for treatment of bilharzia. A decoction of the level up to 750 in altitude. It is said to be a bark and leaves is drunk as a cure for paraly- pioneer species. sis, impotence and convulsions. The leaves are Management Sterculio OPPendiculoto is easy used to treat cerebral malaria. A maceration of to grow from fresh seed the petioles is drunk as a purgative. Trees are Genetic resources and breeding In view planted for shade and as ornamental. of its large geographical distribution Stercu!to Properties The heartwood is pale brown and OPPendiculoto is considered not to be in threat indistinctly demarcated from the sapwood. The of genetic erosion. Intensification of its utilisa- wide growth rings with reddish brown late- tion has been reported to pose a threat locally wood are visible to the naked eye. The wood is in Mozambique. medium-weight, with a density of 580-780 Prospects In view of its potential for plywood kg/ina at 12% moisture content. It is soft and and firewood SterCu!to OPPendiculotO deserves perishable and needs treatment to prevent more research effort. The potential of plant borer damage. The wood is suitable for the parts in treatment of cerebral malaria needs production of wood-cement composites. The verification. seeds contain about 30% oiland 25% protein. Major references Cheek & Dorr, 2007; Botany Deciduous, small to large tree up to Golding, 2002; Roe at a1. , 2002; Ruffo, Birnie & 45 in tall; bole straight, with large buttresses; Tengias, 2002; Wild, 1961. bark surface yellow-white, smooth; young twigs Other references A1berto, Mougel & Zoula- densely rusty brown hairy. Leaves alternate, nan, 2000; All at a1. , 2008; Augustino & Gillah, simple; stlpules early caducous; petiole 5-16 2005; Greenway, 1941; Maingi, 2006; Mensier, cm long; blade ovate, 10-24 cm x 7-20 cm, with 1957; Neuwinger, 2000; Newmark, 2001; 5-7 shallow, acuminate lobes, cordate at base, Paterson & Howland, 1971; Whatsi, 2003 densely scaly hairy when young. Inflorescence Authors C. H. BOSch & D. Louppe an axillary, narrow panicle up to 11 cm long,

. STERCULIA 513

STERCULIAOBLONGAMast. Us$ 9511m' and in 2006 small amounts of veneer at an average price of Us$ 6231 ms. Congo Protologue Onv. ,fl. Trop. Mr. I: 216 (1868) exported about 900 ina of logs per year in 1967- Family Sterculiaceae (APGMalvaceae) 1970, whereas Equatorial Guinea exported Chromosome number2n=36 about 3440 in3 of logs per year in 1996-1998 Synonyms Eribromo oblongo onast. ) Pierre and Gabon on average 3100 ms per year in ex A. Chev. (1917), Sterculio elegontiftoro Hutch. 2001-2005. & Dalzie1(1928). Prdperties The heartwood is pale yellow with Vernacular names Yellow sterculia, white white streaks, and indistinctly demarcated sterculia (En). Byorig (Fr) from the 10-20 cm wide sapwood. The grain Ls Origin and geographic distribution Stercu- straight or slightly interlocked, texture me- lid oblongo occurs in the forest zone from dium to fairly coarse. Quartersawn surfaces Guinea and Liberia east to the Central African have an attractive figure. The wood is oily to Republic and south to Gabon, Congo and the touch and fresh wood has an unpleasant northern DR Congo. sineU Uses The wood (trade names: eyong, okoko, The wood is medium-weight to moderately ohaa) is used in West Africa for flooring, heavy, with a density of 680-840 kg/ina at 12% beams, planks and furniture. It is also suitable moisture content. It air dries slowly, wlth a for construction, veneer, plywood, vehicle bod- tendency for surface checking, distortion and ies, handles, sporting goods, toys, agricultural development of shakes or collapse. The rates of implements, hardboard and particle board. shrinkage from green to oven dry are high: The seeds are eaten. In Cameroon a mixture of 4.5-5.0% radial and 10.2-12.2% tangential. pulped leaves and oilis applied to the chest of Once dry, the wood is usually moderately sta- children as a dressing to cure chest complaints. ble in service In Gabon a bark decoction is drunk as a cure At 12% moisture content, the modulus of rup- for acute stomach-ache. The fibrous bark of ture is 91-153 N/mm2, modulus of elasticity young trees is used as bark cloth for partition- 12,600-17,110 N/mm2, compression parallel to ing huts. grain 49-67 N/min2 shear 3-13 N/min2, cleav- Production and international trade Cam- age 12-18 N/mm and Janka side hardness eroon exported 40,550 in3 of logs in 1997, 4980 N. 29,280 in3 in 2000, 5850 ina in 2003, and 9750 The wood saws well, with moderate blunting ina in 2004. In 2005 it exported about 8000 ina effects on sawteeth. It works wellwith machine of logs at an average price of Us$ 2171m' and tools, but more difficultly with hand tools. A in 2006 about 14,000 ms at Us$ 4851m'. Cam- cutting angle of 20' is recommended in plan- eroon exported 1/20 ms of sawn wood from Trig It finishes with difficulty, and a filler is July 2001 to June 2002, 260 in3 in 2003, 190 in3 needed. Staining and polishing are satisfac- in 2004 and 110 msin 2006. In 2005 it exported tory. Nailing and screwing properties are good, about 1000 in3 of veneer at an average price of but quartersawn surfaces may split on nailing. The wood glues weU. The peeling and slicing properties are good. Steam-bending properties are moderate The wood is not durable, being susceptible to

I attacks by fungi, dry wood borers and termites. . The sapwood is susceptible to Lyetus borers The heartwood is extremely resistant to jin- pregnation with preservatives, the sapwood is permeable. The chemical composition of the ,^ oven-dry wood is: cellulose 36.2-46.2%, furfur- als 11.5-12.4%, pentosans 19.8-21.2%, 11gnin L. 18.2-22.7%, ash 1.3-3.2%. The solubility in hot water is 2.7-6.2%, in alcohol-benzene 1.2-9.2% .. andin I% NaOH 16-23.5%

o Adulterations and substitutes The wood resembles that of Pterogyto and Amphimos spp. Sterculio oblongo - wild Description Deciduous medium-sized to large 514 TIMBERSl

2-2.5 cm x c. I cm, with bright yellow, fleshy seed coat. Seedling with epigealgermination Other botanical information Stercu!to is 3 found throughout the tropics and comprises ^ about 150 species, with about 25 of them in tropical Africa \ Anatomy Wood-anatoimcal description OAWA

;; hardwood codes): \ Growth rings: a growth ring boundaries iridis- ,. , tinct or absent. Vessels: 5: wood diffuse-porous; 13: simple perforation plates; 22: intervessel

, pits alternate; 23: shape of alternate pits po- . Iygona1; 24: intervessel pits minute (S 4 prn); 2 25: intervessel pits small (4-7 prn); 30: vessel- -,, I .:: ray pits with distinct borders; similar to in- 1.1' tervessel pits in size and shape throughout the ' I ':4 ray cell; 42: mean tangential diameter of vessel .. > .. Iumina 100-200 prn; 43: mean tangential di- . ,.. .\ ameter of vessellumina 12 200 I'm; 46::S 5 ves- -" sels per square minimetre. Tracheids and fi- : I. -.,-, * 5:44 ,t. ;\ bres: 61: fibres with simple to minutely bor- - - *-.. dered pits; 66: non-septate fibres present; 70: -...~. .\ I. . fibres very thick-walled. Axial parenchyma: 85 .*t, A ,,,- A axial parenchyma bands more than three cells wide; 87: axial parenchyma Teticulate; 92: four (3-4) cells per parenchyma strand; (93: eight Sterculio oblong0 - I, 60se of 601e, ' 2, flowering (5-8) cells per parenchyma strand). Rays: 98 brunch, . 3, port offrutt larger rays commonly 4- to 10-senate; 99: Iar- Redrown grid udopted by Achingd Sotiri Nur- ger rays commonly > 10'seriate; 102: ray height hornon > I mm; (103: rays of two distinct sizes); 107 body ray cells procumbent with mostly 2-4 tree up to 45 in tall; hole branchless for up to rows of upright and/or square marginal cells; 21 in, up to 150 cm in diameter, straight, cyhn- 110: sheath cells present; 1/4: s 4 rays per min; drical, with buttresses up to 3.5 in high; bark (1/5: 4-12 rays per mm). Stoned structure: surface greyish, with small scales, inner bark 120: axial parenchyma and/or vessel elements fibrous, white to yellow with orange streaks, storied. Mineral inclusions: 136: prismatic crys- strongly resinous, with distinctive smell; crown tals present; 137: prismatic crystals in upright narrow, branches whorled. Leaves alternate, and/or square ray cells; 141: prismatic crystals simple and entire; stipules early caducous; in non-chambered axial parenchyma cells; 142: petiole 2-7 cm long; blade elliptical, 6-15 cm x prismatic crystals in chambered axial paren- 3-8 cm, rounded to obtuse at base, short- chyma cells; 154: more than one crystal of acuminate at apex (3-pointed in saplings), aboutthe same size per cellor chamber. densely stellate hairy below when young, pin- (E. Uetimane, H. Beeckman & P. E. Gasson) nately veined with 10-14 pairs of lateral veins. Growth and development Sterculi0 o610n- Inflorescence an axillary, narrow panicle 3-10 go is classified as a non-pioneer light de- cm long; bracts lanceolate, hairy, c. 3 min long, inarider. This means that seedlings are found caducous. Flowers unisexual, regular, 5- in full shade, but saplings only grow up under merous, cream to greenish-yellow, c. 6 min small gaps in the forest canopy. In Ghana av- long; perianth consisting of free tepals, stellate erage increase in height is about 30 cm during hairy inside; male flowers with c. 10 arithers the first year with 9-year-old trees 3-15 in tall borne on a long common stalk, stalk with long Under the forest canopy in Guinea saplings hairs in basal part; female flowers with ovary were 50 cm tall I year after planting and 125 consisting of 5 carpels united loosely. Fruit cm tall after 2 years, In C6te d'Ivoire trees consisting of (4-)5 woody follicles 10-15 cm showed a mean annual increment in hole di- long, glabrescent, green-yellow to yellow- ameter of 13 mm in the first 14 years after brown, many-seeded. Seeds ovoid, compressed, planting. In the Central African Republic the

. STERCULiA 515 mean annual diameter increment was 1.7-7.3 dry and it has a low natural durability. Over- mm for trees of about 25 cm in hole diameter, exploitation must be avoided as the species Is the fastest growth occurring in an exploited considered vulnerable. Because of its slow and thinned forest. Annual diameterLncrement growth it does not seem to have much potential in secondary forest in Garnero0n averaged 18.6 as a plantation species. min. In this forest individuals of Sterculio ob- Major references Bolza & Keating, 1972; longo were present as emergents wlth an aver- CTRAD Forestry Department, 2003; Farmer, age height of 41 in, whereas the main canopy of 1972; Hall6, 1961; Hawthorne, 1995; Haw- the forest was at 15-30 in. In Ghana flowering thorne & Jongkind, 2006; Irvine, 1961; Laird, occurs in September, October and January and 2000; Normand & Paquis, 1976; Takahashi, fruits are encountered from September to 1978 January. Dispersal of the seeds is probably Other references African Regional Work- done by birds. shop, 1998b; ATIBT, 1986; Bedel at a1. , 1998; Ecology Stercu!to oblongo is most commonly Berti at a1. , 1982; Burki11, 2000; Chudnoff, found in semi-deciduous forest, particularly in 1980; CTFT, 1960b; de in Mensbruge, 1966; de secondary forest. It prefers drier localities Saint-Anbin, 1963; Durand, 1978; Durrieu de Propagation and planting The 1000-seed Madron & Daumerie, 2004; Germain & Barnps, weight is about 1100 g. Seeds have to be sown 1963; InsideWood, undated; Keay, 1958e; Neu- immediately after harvest. Germination begins winger, 2000; 01uwadare, 1998; Parant at al. , after 8-15 days, but the germination rate can 2008; Sallenave, 1964; Vivien & Faure, 1985; be low. In the nursery, the seedlings need to be Wilks & TSSemb6, 2000; Worbes at a1. , 2003 kept in the shade. When they are 25-30 cm tall Sources of illustration Han6, 1961; Wilks they can be planted out under the forest can- & TSSemb6, 2000 opy cover or in forests paths. It is recoin- Authors C. H. BOSch & D. LOUDpe mended to refrain from planting in the fullsun Management In the Central African Repub- 11c, the natural regeneration of Sterculi0 o6- STERCULIAQUINQUELOBA(Garcke)K. Schum. Jongg is good and better in exploited forest than in unexploited forest. In natural forest on Protologue Bot. Jahrb. Syst. 15: 135 (1892) average 9.7 trees per ha were counted in 1995, Family Sterculiaceae (APG: Malvaceae) with a volume of 4.24 ina per ha. The average Vernacular names Egyptian plane tree, Iar- volume of timber of Sterculia oblonga in Gabon ge-leaved star chestnut, large-leaved stercuha was measured in 1963 to be 0.11 ms per ha (En). unalamwezi, inkwera nyani, inukungu, Burning the undergrowth in forest has a nega- inukulamishi, mulende (Sw). tive effect on regeneration of Sterculio oblongo Origin and geographic distribution Stercu- although it seems to be less sensitive to fire lid quinquelobo occurs from eastern DR Congo, than Sterculio rhinopeto!or K. Schum. Burundi and Tanzania south to Zambia, Na- Yield A tree with a diameter of 60 cm yields infoia, Zimbabwe and Mozambique. about 3.3 in3 of log, and trees 100 and 150 cm Uses The wood of Sterculio quinquelobo is in diameter yield 11.6 and 27.8 ms of logs, re- used in Zambia for furniture and mine props spectively In Malawiit is used for coffins and light con- Handling after harvest Freshly harvested struction. It is also suitable for flooring, inte- logs sink in water, so they cannot be trans- nor trim, joinery, turnery, poles, veneer, ply- ported by river. To prevent blue stain and pin- wood and pulpwood. It is also used as fuelwood hole borer attack, logs must be treated with a and for making charcoal. preservative as soon as possible after felling The seeds are edible. Trees are tapped for the and be removed from the forest, sawn and gum, which is traded as 'gum Karaya' together dried rapidly. with the gum of other Sterculio spp. and used Genetic resources The timber of Sterculio as thickener, emulsifier, laxative and denture oblongo is valuable and is selectively exploited. adhesive. The bark is used to make floormats This has led to the species becoming rarer ln and yields fibres that are used to make ropes, Limbe Region, Cameroon and probably else- mats and sacks. Sterculio qutnquelobo is where as well. It is classified as vulnerable in planted as an ornamental and roadside tree. the IUCN Red list, because of habitat decline. A decoction of leaves and bark of thin branches Prospects The wood ofSterculio oblongo has is drunk to cure stomach-ache. A decoction of good strength properties, but it is difficult to the bark from branches is recorded to be given 516 TIMBERSl

as enema as remedy for diarrhoea, whereas tent of the gum of Sterculio quinquelobo, how- boiled bark is reportedIy used as enema ever, ITmits its usefulness in foods and phar- against constipation. A decoction of leaves and maceuticals. The gum of Sterculio quinquelobo roots is drunk to cure malaria. possesses solubility and viscosity properties Production and international trade In Tan. similar to commercial gum arabic from ACocio zania Stercu!to quinquelobo is classified as a spp. , especially In having a rather low viscosity lower value timber species. It is unknown if which is similar in hot and cold water. How- any of its timber or gum is traded in the inter- ever, Sterculio quinquelobo gum does not meet national market. all specifications required of gum arabic and Properties The heartwood is pinkish red much higher amounts of the gum are needed to when freshly cut, darkening to inId-brown on obtain similar results exposure, and distinctly demarcated from the Botany DeciduoussmaUto medium-sized tree, pale yellow, up to 6 cm wide sapwood. The up to 20(-40) in tall; hole branchless for up to grain is straight, texture medium to coarse 4.5 in or more, up to 100 cm in diameter, usu- The numerous very broad rays visible on tan- ally straight and cylindrical; bark surface grey gentialsurfaces give a speckled effect. to pale greenish brown, smooth, flaking, inner The wood is moderately heavy, with a density bark roottled red and fibrous; young twigs with of 690-880 kg/ina at 12% moisture content. Air triangular bud-scales. Leaves alternate, sim- drying is very slow and must be done carefully PIe; stipules early caducous; petiole 6-18 cm to avoid surface checking, splitting, cupping long; blade orbicular in outline, 8-36 cm x 10- and collapse. Boards 2.5 cm thick need 6 34 cm, with 5 triangular, acuminate lobes, months to air dry to 18% moisture content and deeply cordate at base, soft hairy with stellate 2 months more to 14% moisture content hairs on lower surface. Inflorescence an axil- Boards 5 cm thick need 14 months to air dry to Iary, narrow panicle 15-35 cm x 6-11 cm, 25% moisture content, and further drying pro- sticky hairy; bracts 3-7 min long, caducous. gresses only at a rate of I% moisture content Flowers unisexual, regular, 5-merous, pale or per month. Kiln drying is very difficult or jin- yellowish green, c. 3.5 min long; perianth cam- possible. The rates of shrinkage from green to panulate with lobes c. I mm x I mm, outside oven dry are about 3.0% radial and 7.0% tan- sticky hairy; male flowers with 8-10 arithers gential. Once dried, the wood is moderately borne on a common stalk c. 1.5 min long; fe- stable in service. male flowers with ovary consisting of 5 carpels The strength of the wood is rather low. At 12% united loosely. Fruit usually consisting of 5 moisture content, the modulus of rupture is woody follicles 5-8 cm x I-1.5 cm, densely soft 71-75 N/mm2, modulus of elasticity 9500- hairy, yellow-brown, sticky, many-seeded 10,200 N/min2, compression parallel to grain Seeds ellipsoid, 7 mm x 4-7 min, grey-black, 40-42 N/min2, shear 9-11.5 N/mm2 cleavage glabrous. 60 N/mm radial and 59 N/mm tangential, and Sterculio quinquelobo flowers when leaves are Janka side hardness 5960 N. present The wood saws easily and power demand is Sterculio comprises about 150 species and oc- low. Planing and moulding are easy. The wood curs throughout the tropics. In tropical Africa finishes to a good polish, and it bores and drills about 25 species can be found. Sterculio quin- fairly well. Nailing properties are poor; the que!o60 is often confused with Sterculio up- wood tends to split and pre-boring is recoin- pendiculoto K. Schum. The latter species has a mended smooth, non-flaking bark, occurs in more hu- The wood is moderately durable to durable, inid regions, and Is found at altitudes up to 750 being moderately resistant to termites. The in. The differences in flowers and fruits be- sapwood is susceptible to Lyetus borers. The tween the 2 species are distinct. heartwood is extremely resistant to impregna- Ecology Sterculio quinquelobo is found in dry, tion with preservatives, the sapwood is perme- deciduous woodland, often on termite-mounds, able. rocky outcrops and hills, usually in association Carefully harvested and graded gum meets the with Brochystegio and 1806erlinio spp. , up to inolsture content and insoluble matter re- 1650 in altitude. quirements of commercial 'gum Karaya' (a Management Propagation ofSterculio quin- maximum of 200 g/kg and 30 gritg, respec- quelobo can be done by sowing and with cut- tively); it also has low levels of aluminium, tings or truncheons. Seeds can be tested after manganese and cobalt. The high tannin con- collection by submersion in water; those that

. STERcuLiA 517 float are usually damaged by insects and should be discarded. Seeds should be dried to 9-12% moisture content before storage. At a storage temperature of 4'C a germination rate

I of 70% can be maintained for half a year. Ster- . culto quinquelobo tolerates pruning and can be . COPDiced. Star shakes may develop in storage to the fullwidth of the log Genetic resources and breeding Sterculio quinque!o60 is considered not to be at risk of .^ genetic erosion because it is widespread and .. tolerating a range of habitats. However, recent reports indicate that wherever exploitation of .. species with timber of high quality has led to their eradication, trade shifts to Sterculio o quinque!obo timber, resulting in population densities dropping sharply. Prospects Monitoring the utitization of Ster- Sterculio rhinopetolo- wild culto quinquelobo could reveal threats for the species and may indicate potential for sustain- act flatulence. The wood ash yields a vegetable able exploitation. The potential for production salt that is used in cooking and in soap inak- of gum needs to be evaluated. lrig Major references Bolza & Keating, 1972; Production and international trade Cam- Bryce, 1967; Chilufya & Tengnas, 1996; Ruffo, eroon exported about 2000 ina of logs per year Birnie & Tengnas, 2002; Schwartz & Caro, in 2000-2003, 960 in31n 2004 and 1400 ing in 2003. 2006. The same country exported 3725 ms of Other references All at a1. , 2008; Banda at sawn wood in 2003, 2060 main 2004, and 4660 a1. , 2008; Borgerhoff Mulder, Caro & Msago, ms in 2006. In 2003 Gamero0n exported 4000 2007; Chanyenga, 2004; Cheek & Dorr, 2007; in" of plywood, in 2005 about 2000 ms. and in Coates Palgrave, 1983; Mbuna & Mhinzi, 2003; 2006 also 2000 ing. C6te d'Ivoire exported about Minedge & Kaale, 2005; Paterson & Howland, 5000 in3 of logs in 1983, Ghana 3150 in3 of logs 1971; Takahashi, 1978 in 1998 Authors C. H. BOSch & D. Louppe Properties The heartwood is pale to deep reddish brown, and distinctly demarcated from the 4-6 cm wide, white or creamish sapwood STERCULLARHINOPETALAK. Schum. The grain is straight or interlocked, texture medium to coarse. Growth rings are distinct. Protologue Engl. , Monogr. afrik. PIlanzen- Quartersawn surfaces have an attractive fig- Fam. 5: 102 (1900). ure. Family Sterculiaceae (APG: Malvaceae) The wood is moderately heavy, with a density Chromosome number2n=36 of 720-890 kg/ms at 12% moisture content. It Vernacular names Brown sterculia, red air dries very slowly, with serious risk of dis- sterculia (En). tortion and checking. Boards 29 mm thick take Origin and geographic distribution Stercu- about 22 weeks to air dry from green to 18% lid rhinopeto!o occurs from C6te d'Ivolre to moisture content, boards 50 min thick 41 Gamero0n. weeks. To reduce the risk of checking during Uses The wood (trade names: lotofa, wawa- drying, it is recommended to quartersaw the hima) is suitable for construction, flooring, wood. The rates of shrinkage from green to joinery, interior trim, panelling, stairs, high- oven dry are high: 4.9-5.5% radial and 9.5- quality furniture, ship and boat building, tool 11.4% tangential. Once dry, the wood is inod- handles, toys, turnery, poles, veneer and ply- erately stable to unstable in service. wood. The wood is apparently suitable for pa- The wood is tough and hard. At 12% moisture per making but is not as yet used for this pur- content, the modulus of rupture is 1/6-186 pose N/mm2 modulus of elasticity 13,400-18,700 A mixture of the powdered bark and oilis ap- N/mm2, compression parallel to grain 57-81 plied to swellings and taken orally to counter- N/mm2 shear 14-15 N/min2, cleavage 25 518 TIMBERSl

N/mm, Janka side hardness 6180-8050 N and inner bark strongly resinous, red, often with Janka end hardness 5830 N vertical white bands, fibrous; crown narrow, The wood saws and works easily with hand and branches whorled. Leaves alternate, simple machine tools, but tends to blunt tool edges and entire; stipules early caducous; petiole 3- and sawteeth. It finishes well, but a filler is 11 cm long; blade oblong to lanceolate or needed. The wood holds nails and screws well, oblanceolate, 10-30 cm x 4-16 cm, rounded at but pre-boring is recommended to avoid split- base, apex normally obtuse, with short brown ting on nailing. It glues well. The peeling and stellate hairs when young, but glabrescent, slicing properties are fairly good. Steam bend- pinnately veined with 10-14 pairs of lateral Ing properties are moderate veins. Inflorescence an axillary panicle up to 20 The wood is moderately durable. It is moder- cm long, densely hairy. Flowers unisexual, ately resistant to termite attack, but pinhole regular, 5-merous, pale or yellowish green; borer attack sometimes occurs. The sapwood is pedicel c. 5 min long; perianth campanulate susceptible to Lyetus borers. The heartwood is with lobes c. I mm x I min, outside hairy; male extremely resistant to impregnation with pre- flowers with 10 arithers in 2 rows borne on a servatives, the sapwood moderately resistant. short common stalk; female flowers with ovary Adulterations and substitutes The timber consisting of 5 carpels united loosely. Fruit of Sterculio rhinopetolo is often mixed with consisting of I-5 woody follicles 5-7 cm long, that of Herittero utilis (Sprague) Sprague, from many-seeded. Seeds c. 18 min x 8 min, with red which it can be distinguished easily on micro- fleshy seed coat, dangling from the open fruit SCOPic characteristics on white threads. Seedling with epigeal germi- Description Deciduous fairly large tree up to nation 40 in tall; hole branchless for up to 21 in, up to Other botanical information Stercu!to is 120 cm in diameter, straight, cylindrical, with found throughout the tropics and comprises narrow buttresses up to 3 in high; bark surface about 150 species, with about 25 of them in brown, rough-shaggy with rectangular scales, tropical Africa Sterculio foetido L. is a native of tropical Asia It Is a medium-sized tree of up to 30 in tall with digitately compound leaves. In Asia it is considered to produce better quality timber 3 A .. than other Stercu!to spp. It has been intro-

11 duced elsewhere in the tropics, primarily as an I I ornamental. In Ghana, Togo, Benin and Nige- ria it is grown as an ornamental and the seeds are eaten after removal of the seed coat. The seeds contain 24-52% oil. The seeds have a purgative action and eating too many can , cause headache and even abortion. In Kenya and Tanzania it seems to be restricted to bo- -* tanic gardens, but in Mozambique it is widely grown as an ornamental Anatomy Wood-anatointcal description CIAWA

\ hardwood codes):

- .,* Growth rings: 2: growth ring boundaries iridis- tinct or absent. Vessels: 5: wood diffuse-porous; I"'~"-*-L? - ' "* 13: simple perforation plates; 22: intervessel .,. .. .. t, \ .,.,. pits alternate; 25: intervessel pits small(4-7 2 Urn); 30: vessel-ray pits with distinct borders; , "', ~ I ,*'*,.;..*" similar to intervessel pits in size and shape throughout the ray cell; (42: mean tangential .,._",*.,.*".*,, t. **.,..* _,..,. . . diameter of vessellumina 100-200 prn); 43: mean tangential diameter of vessellumina z Stercu!to rhinopet010 - I, 60se of 601e, ' 2, ledj 3, 200 pin; 46: s 5 vessels per square minimetre; port o117uit. 47: 5-20 vessels per square minimetre; 58: Redrown und adopted by AChmod Sotiri Nur- gums and other deposits in heartwood vessels. homerit Tracheids and fibres: 61: fibres with simple to

. STREBLUS 519 minutely bordered pits; 66: non-septate fibres Management In Cameroon the number of present; 70: fibres very thick-walled. Axial pa- trees with a hole diameter over 60 cm ranged renchyma: (76: axial parenchyma diffuse); 78: from 006 to 0.6 per ha and the volume oftim- axial parenchyma scanty paratrachea1; 85: ber from 0.3 to 4.5 matha. In the Mopri forest axial parenchyma bands more than three cells (C6te d'TVoire) 14 Sterculio rhinopetolo trees wide; 92: four (3-4) cells per parenchyma over 10 cm in diameter were counted per ha strand; (93: eight (5-8) ceUs per parenchyma Diseases and pests Stercu!to rhinopetolo Is strand). Rays: 99: larger rays commonly > 10- a host for cotton stainers (Dysdercus spp. ), an senate; 102: ray height > I min; (103: rays of important pestln cotton two distinct sizes); 106: body ray cells procum- Harvesting The minimum feUing diameter bent with one row of upright and/or square is 50 cm in Cameroon and 70 cm in Ghana. marginal cells; 107: body ray cells procumbent Yield In Garnero0n a tree 60 cm in diameter with mostly 2-4 rows of upright and/or square yields about 4.6 in3 of log, and a tree 100 cm in marginal cells; 114: S 4 rays per mm. Stoned diameter 14.3 ms of log. structure: 120: axial parenchyma and/or vessel Handling after harvest The wood is sensi- elements stoned. Mineral inclusions: 136: tive to fungal attack during the air drying prismatic crystals present; 137: prismatic crys- process tals in upright and/or square ray cells; 138: Genetic resources There are no indications prismatic crystals in procumbent ray cells; 140: that Stercu!to rhinopetolo is under threat of prismatic crystals in chambered upright and/or genetic erosion, but in many areas it is rather square ray cells; 142: prismatic crystals in heavily exploited for its timber. Monitoring of chambered axial parenchyma cells; (154: more its exploitation is recommended. than one crystal of about the same slze per cell Prospects The wood of Sterculio rhinopetolo or chamber) has good strength properties, but the tree grows (N. P. MOILel, P. Derienne & E. A. Wheeler) slowly and the wood is difficult to dry and only Growth and development Sterculio rhino- moderately durable. Therefore it does not seem peto!o is classified as a non-pioneer light de- to have much potential as a plantation tree mander. It grows slowly and is shade-tolerant. Major references Bolza & Keating, 1972; Mean annual hole diameter increment in sec- Burki11, 2000; CTRAD Forestry Department, ondary forest in Cameroon averaged 3.6 mm 2003; Farmer, 1972; Hawthorne, 1995; Haw- In this forest some individuals were present in thorne & Jongkind, 2006; Irvine, 1961; Phong- the main canopy, but the majority of individu- phaew, 2003; Takahashi, 1978 als occurred as recruits in the lower storeys. In Other references Agyeman, Swame & C6te d'Ivoire the annual increment in diameter Thornpson, 1999; ATIBT, 1986; Bertault, 1982; was 3.3-7.3 mm in natural forest and 5.0-12.4 Bertault at a1. , 1999; Chudnoff, 1980; GTPT, min in a strongly thinned forest, but trees over 1960a; de in Mensbruge, 1966; Durand, 1978; 40 cm in bole diameter have slower diameter Durand, 1983b; Durrieu de Madron at al. , growth: about 2.5 min per year. In 06te d'Ivoire 1998b; Duviard, 1981; InsideWood, undated; 14-year-old planted trees showed a mean an- Keay, 1958e; Lancaster, 1961; Lemmens, nual growth in diameter of I cm. In C6te Norizo & Sud0, 1995; Normand & Paquis, d'Ivoire and Ghana flowering occurs when the 1976; Parant at a1. , 2008; Riddoch at a1. , 1991; tree is leafless between July and October and Sallenave, 1955; Vivien & Faure, 1985; Worbes fruiting in the dry season. The seeds are dis- at a1. , 2003 persed by birds. Sources of illustration Hawthorne & Jong- Ecology Sterculio rhinopetolo Is found in the kind, 2006; Vivien & Faure, 1985 drier areas of lowland rain forests. Natural Authors C. H. BOSch & D. Louppe regeneration seems good. Under natural conditions, seedlings and saplings can be commonly found in small gaps in the forest, but slightly STREBLUS DIMEPATE (Bureau) C. C. Berg older trees become light demanders Propagation and planting The 1000-seed Protologue Proc. Kon. Ned. Akad. Wetensch. , weight is about 800 g. Multiplication of Sterc!,- C, 91(4): 358 (1988). jig rhtnopetolo using seeds is without problems Family Moraceae with a rapid germination both in light and in Synonyms Pochytrophe dimepote Bureau darkness. Planting in full sun is possible. Ger- (1873), Pochytrophe o60uoto Bureau (1873). mination starts after 4-12 days Origin and geographic distribution Stre- 520 TIMBERSl

difficult and pre-boring is recommended. The wood is very durable, being very resistant to fungal attacks and having good resistance against termites. The heartwood is resistant to

I . impregnation with preservatives, the sapwood is permeable Description Dioecious, evergreen or deciduous shrub or small to medium-sized tree up to 30 in tall, with white latex; branchlets glabres- ,^ cent. Leaves alternate, distichous, simple and entire; stipules fused, 2-6(-9) mm long, L. sparsely to densely hairy; petiole 3-15(-20)

.. min long; blade elliptical to oblong or obovate, (I-)2-IOC16) cm x (0.5-)1.5-6 cm, base acute Q to obtuse, apex acuminate or obtuse, margin often somewhat revolute, leathery, glabrous, pinnately veined with 5-12(-16) pairs of Iat- Streblus dimepote - wild eral veins. Inflorescence a catkin, usually in pairs in leaf axils; male inflorescence 0.5-5 cm 61us dimepote is endemic to Madagascar, where long Including the I-5 mm long peduncle, it is confined to the eastern and north-western many-flowered; female inflorescence 0.5-2.5 cm part of the country long including the 2-15 min long peduncle, 2- Uses The wood (trade name: dipaty) is used 14nowered. Flowers unisexual, sessile, close for posts in house construction because of its together, perianth 1.5-2 min long; male flowers high durability in contact with the soil, and it with 4-parted perianth, membranous, hairy, has been used for railway sleepers. It is not stamens 4, innexed, ovary rudimentary, quad- much used for other purposes because it is dif- rangular, c. 5 min long; female flowers with 4 ficult to work. However, it is suitable for heavy I construction (including bridges and sluice gates), heavy flooring, carpentry, vehicle bod- \ I ;*,., I I 11 ,. Ies, carving, turnery and sliced veneer. The \ ^ wood is also used as fuelwood and for the pro- ...\ I 2 duction of charcoal. The infructescence is edi- I J

,- ble. In traditional medicine an infusion of the . bark and leafis drunk againstjaundice

Properties The heartwood is brown, strongly .~ \ darkening on exposure, and clearly demarcated from the greyish beige sapwood. The grain is straight, sometimes wavy, texture very fine The wood is nicely figured with black veins The wood is heavy, with a density of 800-990 kg/ms at 12% moisture content. It air dries A slowly with a slight risk of distortion. Boards t~

2.5 cm thick take 3-4 months to dry. The rates ~. ^, of shrinkage from green to oven dry are 3.5- 6.4% radial and 6.3-11.5% tangential. The dried wood is not stable in service.

The wood is very hard. At 12% moisture con- L\ tent, the modulus of rupture is 169-236 N/min", modulus of elasticity 15,000-20,000 3 N/min', compression parallel to grain 61-94 N/min', shear 7 N/min2 cleavage 11-17 N/min Styeblus dimepote - I, twig with mole milores- and Chalais-Meudon side hardness 5.3-12.7 eeriees, . 2, port of fernole inflorescence, . 3, twig Due to its hardness, the wood is difficult to cotth inIructescences. work. It usually takes a good polish and fin- Redrown und odopted by AChmod Sotiri Nur- ishes well. It glues easily, but nailing is very hornon

. SWiETENiA 521 free tepals, fairly thick, glabrous to hairy, fined to Madagascar, but it is unclear whether ovary superior, globose to ovoid, style very it is threatened by genetic erosion short, stigmas 2. Fruit a I-seeded drupe 5-8 Prospects The wood of Styeblus dimepote is mm x 6-7 min, with enlarged, succulent, red- beautiful and very durable, but difficult to dish perianth. Seed 4-5 min long work. At present it seems only to be used for Other botanical information Strebluscom- construction, but it may have potential for par- prises about 20 species, and occurs in Africa, quetry, carving and turnery. As such, it may Asia and the Pacific region have export value, but it is difficult to assess Anatomy Wood-anatointcal description CIAWA its potential, because too little information is hardwood codes) available on the availability of wild trees and Growth rings: a growth ring boundaries iridis- the species' potential for domestication tinct or absent. Vessels: 5: wood diffuse-porous; Major references Berg, 1977; Berg, 1988; 13: simple perforation plates; 22: intervessel Gu6neau, Bedel & Thie1, 1970-1975; Salle- pits alternate; 23?: shape of alternate pits po- nave, 1971 Iygona1; 26: intervesselpits medium (7-10 F1m); Other references Gu6neau & Gu6neau, 30: vessel-ray pits with distinct borders; SImi- 1969; InsideWood, undated; Novy, 1997; Per- Iar to intervessel pits in size and shape rier de in Bathie & Leandri, 1952; Schatz, throughout the ray cell; 31: vessel-ray pits with 2001. much reduced borders to apparently simple: Sources of illustration Ferner de in pits rounded or angular; 32: vessel-ray pits Bathie & Leandri, 1952. with much reduced borders to apparently sim- Authors M. Brink PIe: pits horizontal (scalariform, gash-like) to vertical(palisade); 42: mean tangential diameter of vessellumina 100-200 prn; 46: S 5 ves- SunETENtAunCROPHYLLAKing sels per square minimetre; (47: 5-20 vessels per square minimetre); 56: tyloses common Protologue Hook. f. , Icon. PI. 16: t. 1550 (1886) Tracheids and fibres: 61: fibres with simple to Family Menaceae minutely bordered pits; 66: non-septate fibres Chromosome number 2n = 24, 46, 54, 108 present; 69: fibres thin- to thick-walled. Axial Vernacular names Big-leaved mahogany, parenchyma: 85: axial parenchyma bands more large-leaved mahogany, broad-leaved inahog- than three cells wide; (89: axial parenchyma in any, Honduras mahogany (En). ACajou du marginal or in seemingly marginal bands); 93: Honduras, acajou d'Amenque, mahogany eight (5-8) cells per parenchyma strand. Rays: grandes reuilles (Fr). Mogno (Po). 98: larger rays commonly 4- to 10-senate; (103 Origin and geographic distribution Suitet- rays of two distinct sizes); 107: body ray cells enjo macrophyllo is native to the mainland of procumbent with mostly 2-4 rows of upright Central and South America, from Mexico to and/or square marginal cells; (108: body ray Peru, Bolivia and Brazil, which makes it the cells procumbent with over 4 rows of upright most widely distributed Swietenio species. The and/or square marginal cells); 1/4: s 4 rays per mm; 1/5: 4-12 rays per mm. Mineral inclu- SIOnS 136: prismatic crystals present; 137: prismatic crystals in upright and/or square ray

cells; 141: prismatic crystals in non-chambered . axial parenchyma cells ., a '; * (H. Beeckman & P. 06tienne) I. Growth and development Flowering is pro- \ bably year-round, with a peak in September- March. ,^ Ecology Streblus dimepote occurs up to 1000 .. in altitude in dry to humid forest or thickets, often along streams or seashores .@ Management The wood of Styeblus dimepote seems to be obtained from wild trees only, and o no information is available on SIIvicultural management or planting techniques Genetic resources Streblus dimepoteis con- Swigten, o macrophyl!0 -pionted 522 TIMBERSl wood has been internationally traded for over ceeds 120,000 ms. In 1996-2002 the average 400 years. Big-leaved mahogany was intro- annual export of Slutetenio inucrophy!IQ timber duced in India from Belize in 1872, and has was 59,500 in3 from Brazil, 30,500 in3 from since been planted throughout the tropics in Peru, 15,000 ina from Bolivia and 14,000 ms timber plantations and as an ornamental, also from countries in Central America. In 2002 the on a smallscale in tropical Africa. Attempts to price of sawn Swietenio ingcrophyllo timber introduce mahogany to tropical Africa (e. g. in from Bolivia was about Us$ 9801m3. The most Sierra Leone, Nigeria, Uganda) were largely important importer is the United States with unsuccessful due to the severity of shoot borer an import of 76,000 ing/year, which is over 60% attack on young plants. Trial plantations in of global trade. It has been estimated that the Mauritius were initially unsuccessful, but later area planted with Swietenio inocrophyllo in 800ietenio ingcrophyllu was used successfully the tropics is 200,000 ha, butthe area in Africa for reforestation at low elevations. Extensive is insignificant plantations exist in Indonesia, FiJi and Sri Properties The heartwood is reddish or pink- Lanka. ish, the colour darkening with age to a deep red Uses Swietenio wood (mahogany)is regarded or brown, distinctly demarcated from the usu- as the world's finest timber for high-class fur- ally yellowish sapwood, which is up to 40 mm niture and cabinet work. Its popularity is espe- wide. The grain is interlocked, sometimes clany due to its attractive appearance in coin- straight, texture fine to moderately coarse. The bination with ease of working, excellent finish- surfaces are glossy, with golden lustre, and the ing qualities and dimensional stability. It is wood is often nicely figured because of irregu- also often used for Interior trim such as panel- Iar grain. ling, doors and decorative borders. It is used Big-leaved mahogany wood is a medium-weight for boat building, often as a decorative wood for wood. The density is (450,530-670(-840) luxury yachts and ocean liners, but sometimes kg/in3 at 12% moisture content, with that of also as plywood for planking and deck housing. plantation-grown trees often somewhat less Its outstanding technical qualities make it than that of trees from natural forest. The particularly suitable for precision woodwork rates of shrinkage are low, from green to 12% such as models and patterns, Instrument cases, moisture content 1.4% radial and 2.2% tangen- clocks, printer's blocks and parts of musical tial, and from green to oven dry 2.1-3.3% ra- instruments; for these purposes, uniform dial and 2.9-5.7% tangential. The wood sea- straight-grained material is used. Other minor sons well, without much checking or distortion. uses include burial caskets, wood carvings, Planks of 50 min thick can be air dried in 11 novelties, toys and turnery weeks from 40% moisture content to 15%, An oil, which is very bitter and purgative, can boards of 25 min thick in 6 weeks. The wood be extracted from the seeds, but commercial kiln dries satisfactorily when moderate sched- exploitation of the oilseems unlikely. The bark ules are used (temperatures of 43-76'C and is bitter and astringent, and has been used as a corresponding relative humidities of 75-33%). fobrifuge and also for dyeing and tanning Planks of 50 min thick can be kiln dried in ap- leather. A gum is produced for Bombay (India) proximateIy 8 days from 40% moisture content markets from cuts in the bark, where it is sold to 15%, and boards of 25 mm in 4 days. Planks both pure and mixed with other gums. Various of 41 min thick can be kiln dried in 13 days medicinal uses of various parts of the tree are from 70% moisture content to 15%. After dry- reported from tropical America. The crushed ing, the wood is stable in service fruit shells have been used as a potting me- The wood is comparatively soft. At 12% ino1s- dium. Swietenio macrophyllo Is also used in ture content the modulus of rupture is 72-98 reforestation programmes and agroforestry N/min2 modulus of elasticity 9300-12,100 systems, and as a shade tree in young planta- N/mm2, compression parallel to grain 43-62 tions of other timber species, and is occasion- N/min2, shear 13.5 N/mm2 and Janka side ally planted as ornamental hardness 3560 N Production and international trade Maho- The wood saws, planes and moulds easily in gany is one of the most important tropicaltim- both green and dry condition. In general it fin- hers on the world market. Most mahogany ishes to a smooth surface, but a woolly surface traded is from natural stands, and only sinaU may occur on bands of reaction wood or Inter- quantities are available from planted trees. locked grain. Finishing is easy and the wood The annual export from tropical America ex- takes an excellent polish. Gluing and nailing

. SWIETENIA 523 properties are good, but discoloration in contact with iron, copper and brass may occur under .\ . . humid conditions. The wood slices and rotary 6 cuts into fine and decorative veneer, without ;;' I. preliminary treatment, at a peeling angle of '.* * 92'. The veneer can be glued with casein ex- tended with 30% lime, to produce plywood of *' satisfactory quality. Satisfactory results are obtained with pulping (kraft pulp yield of \

49.5%) * The heartwood of trees from natural stands can be reasonably durable, but it is not considered suitable for applications in contact with the ground. Graveyard tests in Indonesia 2 *;!*:'**,,: ,*.\ ,., showed an average service life in contact with ""' ' " ,VPI'.*,. ' .. 4'''4' the ground of 2.7 years. The wood is resistant . to wood-rotting fungi. The sapwood is susceptible to Lyetus borer attack and the heartwood .. ~t, ;#;43t^;;4$;*:-$ may also be attacked by pinhole borers and termites; the wood has little resistance to ina- rine borers. The wood is resistant to Impregna- tion with preservatives by pressure methods, but plantation-grown wood can be amenable to boron diffusion techniques Inn'5 if"""'* The wood contains 39-47% cellulose, 27-31% 11gnin, 16-18% pentosan, 0.5-0.6% ash and Swietenio macrophyllo - I, tree habit, . 2, flower- 0.1% silica. The solubility is 2.4% in alcohol- ing twig, . 3, sectioned mole flower, . 4, sectioned benzene, 0.4% in cold water, 4.5% in hot water female flower, 5, fruit, 6, seed and 19% in a I% NaOH solution. The energy Source. PROSEA value of the wood is 19,600-20,300 kJ/kg. The wood contains essential oil which is rich in smallcymes. Flowers unisexual, but with well- sesquiterpenes. The bark of S"toterrio macro- developed vestiges of the opposite sex, regular, phyllo showed significant in-vivo antimalarial 5-merous, small; pedicel slender, 1.5-2.5 min activity in tests wlth rodents long; calyx with broadly rounded lobes I-1.5 Adulterations and substitutes True ina- mm long, hairy at margins; petals free, slightly hogany timber from Suitetento spp. is hardly contorted in bud, ovate-oblong, 5-6 min long, produced in tropical Africa, but the timber of hairy at margins; stamens united into a tube, several indigenous and more or less related with 10 sessile arithers at the mouth of the species is traded as African mahogany, mainly tube; disk annular; ovary superior, usually 5- Britondrophrogmo and rigyo spp. , which pro- celled, style short, with disk-shaped stigma duce wood with similar characteristics Fruit a woody, elongate-ovoid capsule 10-15(- Description Medium-sized to large monoe- 22) cm long, greyish brown, opening by 5 CIOus tree up to 40(-60) in tall; hole usually valves, many-seeded. Seeds with large wing, straight and cylindrical, branchless for up to flat, 7.5-10 cm long, dark brown, hanging 18(-25) in, up to 150(-200) cm in diameter, down and overlapping in fruit; cotyledons thin often with broad and plank-like buttresses up Seedling with hypogeal germination; first 3 to 5 in high; outer bark of older trees scaly, leaves simple, later ones 3-foliolate or impari- shaggy, deeply longitudinalIy furrowed and pinnate. brownish grey to reddish brown, inner bark Other botanical information Swietenio red-brown or pinkish red; crown dome-shaped, comprises 3 species and is most closely related consisting of a few large, ascending branches to Khoyo, which differs in its more globose Leaves alternate, panpinnate with (2-)3-6(-8) fruits and seeds narrowly winged allround the pairs of leaflets; stipules absent; leaflets OPPo- margin. The 3 Spyietenio species are difficult to site, ovate-elliptical, 8-18 cm x 3-5.5 cm, en- distinguish from each other. SUIietenio macro- tire, glabrous, pinnately veined. Inflorescence phyl!o differs from Swietenio inohogoni(L. ) an axillary panicle 10-20 cm long, consisting of Jacq. in its larger leaflets, fruits and seeds, and 524 TIMBERSl from Swieteniu humilis Zucc. in its stalked and ground with a position oblique to the main shortly acuminate leaflets (sessile and long- stem. However, an unbranched stern up to 10 acuminate in Swietenio humtlis) and dark in tall may be formed before branching. Initial brown seeds male brown in Suietenio humilis) growth is fast, depending on site conditions. The natural distribution areas of the three Under optimal conditions, seedlings may reach species show only very slight overlap, but in 3 in in one year and 6 in in two years, The av- areas where 2 species occur hybrids may be erage height of 2-year-old seedlings in experi- found. mental plantations in Nigeria at the beginning Swietenio humilis has also been introduced in of the 1960s was 2 in, but there is no informa- a few places in tropical Africa (e. g. in Malawi). tion on growth after 2 years. The seedlings and It is a small to medium-sized tree up to 20 in saplings are intoIerant of deep shade although tall, originating from Central America. Its tim- they will survive. Trees planted in the open her is of little economic importance. field under optimal conditions may occasionally Anatomy Wood-anatorritcal description onWA reach 50 cm in diameter in 25 years. In forest hardwood codes) in the natural distribution area, trees take on Growth rings: (I: growth ring boundaries dis- average 105 years to reach a bole diameter of tinct); (2: growth ring boundaries indistinct or 80 cm, but the average diameter growth in absent). Vessels: 5: wood diffuse-porous; 13: natural forest has also been estimated at 0.4 simple perforation plates; 22: intervessel pits cm/year. In well-managed stands it may be up alternate; (23: shape of alternate pits polygo- to 1.4 cm/year. The maximum increment in na1); 24: intervessel pits minute (S 4 F1m); 30: height occurs at an age of 5-15 years, in di- vessel-ray pits with distinct borders; similar to ameter at 5-10 years and in timber volume at intervessel pits in size and shape throughout 20-25 years. Buttresses begin to develop soon, the ray cell; 42: mean tangential diameter of often already when the hole diameter is only 10 vessellumina 100-200 pin; (46: S 5 vessels per cm. Trees of 200 years old with a log diameter square minimetre); 47: 5-20 vessels per square up to 190 cm and timber volume up to 29.5 ms minimetre; 58: gums and other deposits in have been recorded. heartwood vessels. Tracheids and fibres: 61: Under favourable conditions, young trees may fibres with simple to minutely bordered pits; start to flower and produce seed when 12 years 65: septate fibres present; 66: non-septate fi- old, rarely already when 8 years old, but under bres present; 69: fibres thin- to thick-walled. natural conditions profuse production of seed is Axial parenchyma: 78: axial parenchyma much later. Flowering trees have about 10 scanty paratrachea1; 79: axial parenchyma times as many male as female flowers, but the vasicentric; 89: axial parenchyma in marginal flowers of both sexes are very similar. Trees or in seemingly marginal bands; 93: eight (5-8) are sometimes functionally dioecious. In mixed ceUs per parenchyma strand; 94: over eight inflorescences, male flowers open first, but self- ceUs per parenchyma strand. Rays: 97: ray pollination may occur. The flowers are proba- width I-3 cells;(98:larger rays commonly 4- to bly pollinated by bees and moths, and outcross- 10'seriate); 106: body ray cells procumbent ing is the rule. Fruits may be produced once a with one row of upright and/or square marginal year, and mast production of seeds occurs Ir- cells; 115: 4-12 rays per mm. Stoned structure regularly. Mature fruits can be found 10-11 (1/8: all rays stoned); (122: rays and/or axial months after flowering. The seeds are provided elements irregularly stoned). Secretory ele- with a thin tail-like wing that makes them merits and cambial variants: 131: intercellular rotate when they fall, and are thus dispersed canals of traumatic origin. Mineral inclusions by wind occasionally as far as 500 in from the (136: prismatic crystals present); (137: pris- mother tree. matic crystals in upright and/or square ray Ecology Under natural conditions big-leaved cells); (141: prismatic crystals In non- mahogany thrives in both deciduous and ever- chambered axial parenchyma cells); (142: green rain forest and occurs scattered or in prismatic crystals in chambered axial paren- small groups, but more than 4-8 mature trees chyma cells). per ha are rarely encountered. It has been (E. Uetimane, H. Beeckman & P. E. Gasson) claimed that under natural conditions it regen- Growth and development Healthy seeds erates in essentially even-aged stands after start germinating about 10 days after sowing. catastrophic disturbances of the forest, e. g. Young trees have straight and slender stems, hurricanes, fire and flooding. Adult mahogany and branches are formed 2-3 in above the trees may survive such events because of their

. SWiETENiA 525 strong buttresses and resistance to fire, and ing in the field is usually 2.5-3 in x 2.5-3 in spread their seeds into the resulting gaps or Profuse natural regeneration has been found in clearings. However, it has also been demon- many mature mahogany plantations, and this strated that big-leaved mahogany does not may be used to establish the next crop, reduc- require gaps for seed germination, probably ing nursery and plantation establishment except in evergreen forest where little light costs penetrates through the canopy Management Although natural regeneration The optimum annual rainfallis 1400-2500(- in Swigtenio inocrophyllo stands may be PIen- 3500) min with a dry period of 0-4 months. tiful, it is usual to plant seedlings raised in Swieteniu inocrophy!IQ grows from sea-level to nurseries. One-year-old seedlings attain opti- 1500 in altitude, in areas with a mean annual inal height and diameter growth when rertil- temperature of 20-28'C, the range of the cold- ized with 3.6 g N, 2.4 g P205 and 3.6 g K20 est and warmest month being 11-22'C and 22- Phosphorus appears to be the most limiting 30'C, respectively. It is largely unspecific as to element for seedling growth. soil requirements. Within its natural range It Monoculture plantations of mahogany are sus- has been found on alluvial soils, volcanic soils, ceptible to pests, and for that reason mixed heavy clays, latentic soils, soils derived from plantations with other fast-growlrig species are limestone, granite and other sedimentary, ig- often preferred. Thinning usually starts 6 neous or metamorphic rock formations and years after planting, and progressiveIy reduces even on shallow rendzinas. In plantations it the number of trees to 220-400 treesA1a in performs best on deep, fertile, well-drained plantations of 20 years old, and to 120-150 soils with a pH of 6.5-7.5. It does not tolerate treesA1a in 35-year-old plantations. The rota- waterlogging. In tropical milerica mahogany Is tion is usually 40-60 years' among the pioneer species reoccupying de- Diseases and pests Several fungal diseases graded agricultural land. It has been shown affect seedlings in tropical Asia and islands of that teak is outcompeted by mahogany in a the Pacific, but these rarely cause serious mixed stand damage. Shoot-boring moths of the genus Hyp- Propagation and planting Fruits must be sipylo are the major limitation to plantation collected when ripe to achieve a good germina- establishment, and this is also the case in tion rate. The fruits open after 2 days of stor- tropical Africa, where Hypsjpylo robusto is also age and the seeds are very viable with a ger- a major pest in Britondrophrogmo and maya mination rate of 60-95%. The seeds can be plantations. Attack by Hypsipylo can be suc- stored up to 2 months, or longer (up to one cessfully controlled with insecticides under year) if kept cool (2-5'C) in sealed containers nursery conditions. funbrosia beetles may at- at about 45% relative humidity. The weight of tack the wood, creating 'pin-holes' visible in 1000 seeds is 400-500 g. sawn timber. Termites may also cause damage Seeds are sown in the nursery in drills of 2-4 in plantations. cm deep, or they are pushed into flat beds, Harvesting Mahogany plantations are clear- leaving part of the wing exposed. Seed germi- cut when the rotation age (40-60 years) or a log nation starts after about 10 days and continues diameter of 50 cm have been reached, and sub- for about 20 days. The seed bed should be well sequently replanted with nursery-raised seed- manured and shaded, but after 3-4 weeks the lings seedlings grow best in full sunlight. Seedlings Yield In rotations of 50-60 years, average may have a 70% survival rate, even if planted annual volume increments of 15-20 mama can during the dry season, provided they are par- be achieved for plantations, on poor sites 7-11 tiaUy shaded and watered whenever soil ino1s- in3ftja. ture drops below 30%. For field planting, bare- Handling after harvest Mahoganylogsfloat root stock, balled seedlings, stumps (stem in water and can be transported by river. The length 20 cm, root length 20-40 cm, diameter sapwood is susceptible to staining; an anti- of root collar 0.5-2.5 cm) or striplings are used sapstain dip should be used during drying Batted seedlings of 3-4 months old are pre- Genetic resources Populations of mahogany forred. Cuttings are relatively easy to establish have been depleted through centuries of coin- from 3-year-old plants, but more difficult later mercial exploitation in Central and South Suiietenio macrophyl!o can be propagated by America. Although it is widespread, Suitetento in-vitro culture and grafting; the latter method inderophy!IQ has become rare in parts of its has been used to establish seed orchards. Spac- natural area of distribution. It has been in- 526 TIMBERSl cluded in Appendix U of CITES since 2003, in at a1. , 1996; InsideWood, undated; Lamb, 1966; addition to the other two S"totenio species, Lowe at a1. , 2003; Richter & Danwitz, 2000; which were already included. This makes it SnOok, 1996; Styles, 1981; Tinier, 1995; subject to strict regulations concerning export Weaver & Francis, 1988; Ya0, 1981. and trade; exporting countries are required to Sources of illustration Prawirohatmadjo verify that each shipment was legally acquired at a1. , 1993 and that the harvest was non-detrimental to Authors R. H. M. J. Lemmens the survival of mahogany. In Brazil Swietenio Based on PROSEA 5(I): Timber trees: Major mererophyllo has been exploited only for the commercial timbers. national market since 2003, and the minimum diameter limit for exploitation has been raised from 40 cm to 60 cm. There is no restriction for SunETENIAMAHAGONl(L. ) Jacq trade and export of plantation-grown timber. In tests in Central America using seedlings Protologue Enum. syst. PI. : 20 (1760). raised from seed of different origin, significant Family Menaceae variation has been found both within and be- Chromosome number 2n= 12.18, 24, 36, 42, tween Swieteniu ingcrophyllo populations, par- 46, 48, 54, 56, 60, 108 ticularly in terms of height growth and suscep- Vernacular names SinaU-leaved mahogany, tibility to pest attack. An analysis of the ge- narrow-leaved mahogany, West lndian inahog- netic diversity in and between populations in any, Spanish mahogany, Cuban mahogany, Brazil showed high variation. However, an Caribbean inahagoni (En). ACajou Saint- analysis of the genetic diversity in Costa Rica Domingue, acajou de Cuba, mahogany petites indicated very little gene flow reuilles (Fr). Breeding Selection work has been carried out Origin and geographic distribution Swiet- on an experimental scale (e. g. in Indonesia), enjo inohogoniis native to southern Florida especially towards Improving growth and ger- and the islands of the Caribbean Sea. The wood ruination rates. Artificial hybridization be- has been internationally traded for over 400 tween Suiteterrier spp. is possible, and hybrids years. SinaU-leaved mahogany was introduced often show promising features, combining for in India from the West Indies in 1795, and instance the fast growth of Swietertio inocro- since then has been planted throughout the phyllo and the superior-quality wood of Swiet- tropics on a small scale in timber plantations enjo inghogoni, and having greater resistance and as an ornamental, occasionally also in to diseases and pests tropical Africa. Attempts to introduce small- Prospects A major problem in mahogany leaved mahogany to tropical African countries plantations is the susceptibility to Hypsipylo (e. g. Nigeria, Kenya, Uganda and Tanzania) attack. Research priority should be given to the were largely unsuccessful due to the severity of selection of resistant trees which are fast- shoot borer attack on young plants. It has been growing and have an acceptable wood quality planted more successfully in Reunion and The establishment of optimal methods of vege- Mauritius, and small-scale plantations exist in tative propagation is urgently needed. In-vitro Egypt. It is occasionally planted in gardens, micropropagation techniques have already e. g. in Senegal and Mozambique been developed, and these offer possibilities for Uses Suitetento wood (mahogany) is regard- the multiplication of pest-resistant genotypes. ed as the world's finest timber for high-class Developments in research and breeding of ina- furniture and cabinet work. Its popularity is hogany should be closely monitored because especially due to its attractive appearance in they may offer new opportunities for the estab- combination with ease of working, excellent lishment of mahogany plantations in tropical finishing qualities and dimensional stability. It Africa, in addition to African mahogany (Err- is also often used for interior trim such as pan- tondrophrogmo and nayo) and also in the elhng, doors and decorative borders. It is used light of the successful Swietenio plantations in for boat building, often as a decorative wood for tropical Asia and islands of the Pacific Ocean luxury yachts and ocean liners, but sometimes Major references Blunde11, 2004; Brown, also as plywood for planking and deck housing Jennings & Clements, 2003; ITT0, 2004; Its outstanding technical qualities make it Mayhew & Newton, 1998; Prawirohatmadjo at particularly suitable for precislon woodwork a1. , 1993; WorldAgroforestry Centre, undated. such as models and patterns, instrument cases, Other references Burki11, 1997; HeIgason clocks, printer's blocks and parts of musical

. SWIETENIA 527 instruments; for these purposes, uniform preliminary treatment, at a peeling angle of straight-grained material is used. Other minor 92* uses include burial caskets, wood carvings, The heartwood of trees from naturalstands Is novelties, toys and turnery durable, but is not considered suitable for ap- An oil can be extracted from the seed kernels PIications in contact with the ground. Grave- which might be of some commercial value. The yard tests in Indonesia showed an average bark has been used for dyeing and tanning service life in contact with the ground of 3.3 leather. Various medicinal uses of various years. The wood is resistant to most, but not parts of the tree are reported from tropical all, wood-rotting fungi, and liable to termite America. The crushed fruit shells have been attack. used as a potting medium. Suitetento inohogoni Methanol extracts of Swietenio inghogoni bark is also used in reforestation programmes and showed inhibitory activity on HIV-I protease as a shade tree in young plantations of other Ether extracts of seeds inhibited platelet ag- timber species, and is occasionally planted as gregation, probably due to the presence of an ornamental. tetranortrlterpenoids. Production and international trade Suiet- Botany Medium-sized tree up to 30 in tall; enjo inghogoniis no longer traded coinmer- hole often short and much-branched, up to 100 cially as a result of intense over-harvest in the cm in diameter, often with short and blunt past. Only smallquantities of plantation-grown buttresses; outer bark of older trees scaly and timber are occasionally available on the inter- reddish brown; crown rounded. Leaves alter- national market, in which Africa plays no role. nate, panpinnate with 2-4(-5) pairs of leaflets; Properties Small-leaved mahogany wood is stipules absent; leaflets opposite, ovate- a medium-weight hardwood, which is coinpara- lanceolate to ovate-elliptical, (4-)5-6(-8) cm x tively soft. The heartwood is reddish or pinkish, (1.5-)2.5-3.5 cm, entire, glabrous, pinnately the colour darkening with age to a deep red or veined. Inflorescence an axillary panicle (5-)8- brown, distinctly demarcated from the usually 15(-18) cm long, consisting of small cymes yellowish sapwood, which is up to 40 min wide. Flowers unisexual, but with well-developed The density of the wood is 560-850 kg/ina at vestiges of the opposite sex, regular, 5-merous, 15% moisture content, with that of plantation- small; pedicel slender, 1.5-3 mm long; calyx grown trees often somewhat less than that of with broadly rounded lobes I-1.5 min long, trees from the forest in the natural area of dis- glabrous; petals free, slightly contorted in bud, tribution. The grain is interlocked, sometimes ovate-oblong, 3-4.5 mm long, glabrous; sta- straight, texture fine to moderately coarse. The mens united into a tube, with 10 sessile an- surfaces are glossy, and the wood is often thers at the mouth of the tube; disk annular; nicely figured because of the irregular grain. ovary superior, usually 5-celled, style short, At 15% moisture content the modulus of rup- with disk-shaped stigma. Fruit a woody, ovoid ture is 54.5 N/min2, modulus of elasticity 9555 to almost globose capsule (4.5-)6-10 cm long, N/min2, compression paraUel to grain 37 dark brown or blackish, opening by 5 valves, N/mm2 shear 7-7.5 N/min2, cleavage 55 N/min many-seeded. Seeds winged, flat, (2-)4-5(-6) radial and 63.5 N/min tangential, Janka side cm long, dark brown, hanging down and over- hardness 3840 N and Janka end hardness 3840 lapping in fruit. Seedhng with hypogeal germi- N. The rates of shrinkage are low. The wood nation; cotyledons thin; first leaves simple, seasons well, without much checking or distor- later ones 3-foliolate or imparipinnate before tion and kiln dries satisfactorily when moder- becoinlrig panplrinate. ate schedules are used. After drying, the wood Scoretenio comprises 3 species and Is most is stable in service closely related to Khoyo, which differs in more The wood saws, planes and moulds easily in globose fruits and seeds which are narrowly both green and dry condition. In general it fin- winged all round the margin. The 3 Swietenio ishes to a smooth surface, but a woolly surface species are difficult to distinguish from each may occur on bands of reaction wood or inter- other. Swietenio inghogoni differs from Swiet- locked grain. Finishing is easy and the wood enjo macrophy!10 King in its smaller leaflets, takes an excellent polish. Gluing and nailing fruits and seeds, and from Suretenio humi!is properties are good, but discoloration in contact Zucc. in its stalked and shortly acuminate leaf- with Iron, copper and brass may occur under lets (sessile and long-acuminate in Suitetento humid conditions. The wood slices and rotary humi!is) and dark brown seeds (pale brown in cuts into fine and decorative veneer, without Slutetenio humilis). The natural distribution 528 TIMBERSl areas of the three species show only very slight moisture drops below 30%. For field planting, overlap, but in areas where 2 species occur bare-root stock, balled seedlings, stumps (stern hybrids may be found. length 20 cm, root length 20-40 cm, diameter Healthy seeds start germinating about 15 days of root collar 0.5-2.5 cm) or striplings are used. after sowing. Young trees have straight and Cuttings are relatively easy to establish from slender stems, and branches are formed 2-3 in 3-year-old plants, but more difficult later above the ground with a position oblique to the Spacing in the field is usually 2-3 in x 2-3 in main stem. Initial growth is fast, depending on In Egypt spacings o12 in x 2 in have been used; site conditions. The seedlings and saplings are wider spacing (3 in x 3 in) had no effect on bole intoIerant of deep shade and exhibit their fast- diameter growth. Profuse natural regeneration est growth with overhead light and lateral has been found under many mature mahogany shading. The average diameter growth in per- plantations, and this may be used to establish manent plots in Cuba was 0.55 cm/year. the next crop, reducing nursery and plantation The tree is evergreen in its natural habitat, but establishment costs. However, it is usual to may lose its leaves during the dry season in plant seedlings raised in nurseries seasonal climates. Flowering trees have about Monoculture plantations of mahogany are sus- 10 times as many male as female flowers, ceptible to pests, and for that reason mixed which are very similar. Trees are sometimes plantations with other fast-growing species are functionally dioecious. In mixed inflorescences, often preferred. Early pruning is needed for a male flowers open first, but self-pollination proper hole form. Thinning usually starts 6 may occur. The flowers are pollinated by bees, years after planting, and progressiveIy reduces moths and thrips. Fruits may be produced once the number of trees to 220-400 treesA1a in a year, but mast production of seeds occurs plantations of 20 years old, and to 120-150 irregularly. Mature fruits can be found 8-10 trees/}Ia in 35-year-old plantations. The rota- months after flowering. The seeds are provided tion is usually 40-60 years. with a thin tail-like wing that makes them Several fungal diseases affecting seedlings rotate when they fall, and are thus dispersed have been reported in tropical Asia and islands by wind asfar as 500 in from the mother tree of the Pacific, but these rarely cause serious Ecology Under natural conditions small- damage. Shoot-boring moths of the genus Hyp- leaved mahogany thrives in both dry and moist sipylo are the major limitation to plantation forest, often on limestone, up to 800 in altitude. establishment, and this is also the case in The optimum annual rainfallis 1000-1500(- tropical Africa, where Hypsjpylo robusto is also 2500) mm with a dry period of 0-4 months a major pest in Britondrophrogmo and Maya 800ietenio inghogoni grows in areas with a plantations. In Mauritius young Swigtenio mean annual temperature of 15-32'C, with inahogonitrees are also attacked by this pest. 11-12'C in the coldest month. It is largely un- Under nursery conditions attack by Hypsipylo specific as to soil requirements, but grows best can be successfully controlled with insecticides on well-drained, rich but sandy soils Ambrosia beetles may attack the wood, creat- Management Ripe fruits must be collected ing 'pin-holes' visible in sawn timber. Termites to achieve a good germination rate of seeds may also cause damage in plantations. The fruits open after 2 days of storage and the Mahogany plantations are clear-cut when the seeds have a germination rate of 60-95%. The rotation age has been reached (40-60 years), seeds can be stored up to 2 months at ambient and subsequently replanted with nursery- temperatures, or up to one year if kept cool(2- raised seedlings. Average annual volume in- 5'C) in sealed containers at about 45% relative crements of Swietenio inohogoni plantations humidity. are usually slightly lower than those of S"jet- Seeds are sown in the nursery in drills of 2-4 enjo macrophyllo. The logs float in water and cm deep, or they are pushed into flat beds, can be transported by river. The sapwood is leaving part of the wing exposed. Seed germi- susceptible to staining; an anti-sapstain dip nation starts after about 10 days and continues should be used during drying. for about 20 days. The seed bed should be well Genetic resources and breeding Natural manured and shaded, but after 3-4 weeks the populations of small-leaved mahogany have seedlings grow best with only lateral shading. been depleted through centuries of commercial Seedlings may have a 70% survival rate, even exploitation. It is considered to represent a ifplanted during the dry season, provided they prime example of genetic erosion resulting are partially shaded and watered whenever soil from past exploitation of preferred genotypes

. SnipHONiA 529

(long, straight holes), with populations remain- phonio g!obu!ifero has a remarkable distribu- ing almost entirely composed of highly tion, occurring naturally in the rainforest of branched trees. It has been included in Appen- tropical America and Africa. In tropical Amer- dix H of CITES since 1992, which makes it ica it occurs from Mexico to Brazil and Peru subject to strict regulations concernlrig export and in tropical Africa from Guinea Bissau to and trade. There is no restriction for trade and Tanzania, western Zambia and Angola, possi- export of plantation-grown timber bly also in Madagascar. Marine dispersal of Prospects A major problem in mahogany whole trunks has been postulated as the most plantations is the susceptibility to Hypsipylo plausible cause of the present distribution. It is attack, Suretenio inohogoni often being even supported by the observations that it is coin- more susceptible than Swietento inderophyllo. mon along rivers and propagates vegetativeIy The latter species or hybrids between the two in open habitats. Distribution through disper- species seem to be more promising for planta- sal of seed by sea is unlikely as the seeds do tion establishment, also because of higher not survive desiccation or prolonged immersion growth rates, in addition to African mahogany in seawater. Seeds also pass through the diges- (Britondrophrogmo andKhoyo) tive tracts of animals too fast to allow them to Major references Maybew & Newton, cross the Atlantic Ocean. 1998; Prawirohatmadjo at a1. , 1993; World Uses The wood ofSymphonioglobulifero called Agroforestry Centre, undated 'manil','manni','DSSol' (Gabon) or 'boarwood' in Other references Burki11, 1997; CSIR, 1976; international trade, is a general purpose tim- CTFT, 1993; Gilman & Watson, 1994; HeI- her used in construction, floorlrig and carpen- gason at a1. , 1996; Lamb, 1966; Matsuse at al. , try and for railway sleepers, boxes, crates, coo- 1997; Styles, 1981 perage and sport articles, and traditionally to Authors R. H. M. J. Lemmens make toolhandles and canoes. It is suitable for Based on PROSEA 5(I): Timber trees: Major making plywood commercial timbers. The bark gum, called 'hog gum , manL wax or 'karamani wax', is Insoluble in water and serves as glue, e. g. for joinlrig wood, fixing tool SnipHONiAGLOBULiFERAL. f handles, and to caulk boats and calabashes. It is also used to make torches and candles. In Protologue Suppl. PI. : 302 (1782) Guyana an ammonia-soluble dye is prepared Family Clusiaceae (Guttiferae) from it, which is used to give leather a rich Synonyms Symphonio gobonensis oresque) brown colour. In Ghana a tomc prepared from Pierre (1896), the bark is taken as appetizer and stomachic, Vernacular names Boarwood, hoggum, chew and an extract of the bark is used to treat river stick (En). Manil marecage, mani(Fr). Mundela, blindness. Boiled bark and roots are used as a 61eo bar50 (Po). Mziwaziwa (Sw). wash to treat itch, and the resin Is used to Origin and geographic distribution Sym- treat wounds and prevent skin infections. The bark and heartwood are used in Garnero0n as laxative for pregnant women and as a general tonic. In Gabon the bark is used as an emetlc to treat chest complaints and the gum Ls applied against scabies. In Uganda the bark is . used to treat cough in children. The gum Is taken internally in Nigeria to treat gonorrhoea \a 96 and as a diuretic. In DR Congo sap from the 39 ^ leaves is sniffed up the nose to stop it bleeding.

. ,A Because of its attractive red flowers Sympho- itto globu!ifero may be grown as an ornamental \.. tree a Production and international trade Most .. Symphonio globulifero timber in international

o trade originates from tropical America; quantities exported from Africa are much smaller. No import/export statistics are available. Some- Symphonioglobulifero - wild times the timber is traded mixed with that of 530 TIMBERSl

Angeno spp 4 ..* Properties The heartwood is buff-brown with shades of yellow, rose or orange, and distinctly demarcated from the grey-yellow, 2-5 cm wide I 3 sapwood. The grain is generally straight, sometimes interlocked, texture medium to coarse The wood has medium lustre and a mealy appearance, with conspicuous lines and arches on *, . ^ -, the radial surface and inottling on the tangen- . tial surface. The wood is classified as medium-weight to moderately heavy. The density is 530-720(- I; 750) kg/ms at 12% moisture content. The ^,-=-

^ shrinkage rates are high, from green to oven dry 4.36.6% radial, 9.7-10.7% tangential. The ~' .,..,"V.,.*,'"?*A# *.. wood dries moderately rapidly, with high risk I of distortion and checking. Air drying under I .*' cover and end coating are recommended. Once dry, the wood is poorly stable in service At 12% moisture content, modulus of rupture is ,

82-181 N/min2, modulus of elasticity 12,500- ., . I ,. 18,400 N/mm2, compression parallel to grain :: 42-67 N/min', shear 7-10 N/mm' cleavage 15- 17 N/mm, Janka side hardness 4400-5040 N 2 and Janka end hardness 5640 N Workability is good with both hand and ina- Symphonioglobuli/eru - I, base o160!e, 2, nower- chine tools; ordinary sawteeth and cutting tools ing twig, 3, fruit, . 4, seed. can be used. The nail- and screw-holding ca- Redrown ond udopted by AChmud Sotiri Nur- pacity Is good, but pre-boring is necessary. The hornon wood can be glued well with most glues. It can be rotary-cut for veneer and is suitable for ex- absent; petiole 0.5-2 cm long, channelled tenor and interior layers of plywood. The wood above, finely transversely wrinkled; blade ellip- Is moderately durable, but should not be used tical to lanceolate, 5-12.5 cm x I-5 cm, base In contact with the ground under prolonged curieate, apex obtusely acuminate, leathery, wet conditions. It Is susceptible to termite at- mostly dark glossy green, with closely parallel tack. Treating with preservatives is difficult secondary venation. Inflorescence a sessile, The wood contains 47-52% cellulose, 24% 11g- umbel-like, many-flowered cyme, terminal on rim and 17-20% pentosan. The energy value of short lateral branches. Flowers bisexual, regu- dry wood Is 18,450 kJ/kg Iar, 5-merous; pedice1 0.5-2.5 cm long, longer Poly-ISOprenylated herizophenone derivatives in fruit; sepals circular, kidney-shaped or (guttiferones A-D) have been isolated from the ovate, 2-5 mm x 2.5-7 min; petals circular, roots. These compounds showed HTV-inhibitory 0.5-1.5 cm x 1.5-2 cm, crimson or scarlet, effects in cell cultures. The root bark contains waxy; disk cupule-like, 5-angled, 1.5-4 mm prenylated xanthones (globulixanthones A-E), thick, margin entire or undulate, persistent; which showed cytotoxicity in cancer celllines. stamens in 5 groups of 3-4, basalIy merged Description Evergreen medium-sized tree up Into a 3.5-10 min long tube, with an outer to 25(-40) in tall, with sticky, yellowish latex, whorl of stainlnodes; ovary superior, ovoid, 5- glabrous; hole straight, cylindrical, branchless celled, styles 5, fused at base, spreading or for up to 21 in, up to 80(-100) cm in diameter, recurved. Fruit a broadly ellipsoid or globose without buttresses but with stilt roots or berry 1.5-4.5 cm x 2-3.5 cm, finely warty, with pneumatophores in swampy localities; bark copious yellow fluid turning brown on expo- buff to greenish yellow or grey-brown, smooth sure, I-3-seeded. Seeds compressed ovoid, 1.5- or vertically fissured or with lenticels in verti- 2 cm x I-1.5 cm, testa tin, marbled. Seedling cal rows, 5-8(-15) mm thick; crown rounded, with hypogeal germination and thickened hy- with numerous horizontal, opposite branches. pocotyl Leaves opposite, simple and entire; stipules Other botanical information Symphonio

. SnipHONiA 531 comprises about 20 species, all except Sympho- fruits. In regularly flooded locations Sympho- itto globulifero confined to Madagascar itto globulifero often develops stilt roots and Anatomy Wood-anatomical description OAWA pneumatophores hardwood codes): Ecology Symphonio globulifero is a specles Growth rings: 2: growth ring boundaries Iridis- of evergreen mixed humid forest or freshwater tinct or absent. Vessels: 5: wood diffuse-porous; swamp forest with an average rainfall of 650- 13: simple perforation plates; 22: intervessel 2100 min and 23-27'C mean annual tempera- pits alternate; (23: shape of alternate pits po- ture. It occurs from sea-level up to 2600 in alti- Iygona1); (26: intervessel pits medium (7-10 tude in East Africa. It also occurs at Inner pin)); 27: intervessel pits large (^ 10 prn); (30: edges of mangroves that are only occasionally vessel-ray pits with distinct borders; similar to flooded with salt water. intervessel pits in size and shape throughout Propagation and planting Fresh seed ger- the ray cell); 31: vessel-ray pits with much re- inmates best at 25-30'C, under natural condi- duced borders to apparently simple: pits tions within one month, but it dies quickly rounded or angular; 43: mean tangential di- upon dehydration. All seeds die below 12*C. ameter of vessellumina Z 200 pin; 46: S 5 ves- Germinated seeds can be stored at 15'C under sels per square minimetre; (47: 5-20 vessels humid conditions for about a year and seed- per square minimetre); 56: tyloses common; tings kept at this temperature develop slowly (58: gums and other deposits in heartwood ves- but resume normal growth when transferred to sels). Tracheids and fibres: 61: fibres with SIm- 25'C. Seed requires shade for germination and PIe to minutely bordered pits; 66: non-septate in closed forest propagation is by seed only; in fibres present; 70: fibres very thick-walled. open areas propagation Is exclusively by root Axial parenchyma: 80: axial parenchyma all- suckers. In Gabon it was found that natural form; 82: axial parenchyma winged-allform; 83: regeneration in undisturbed forest is good, but axial parenchyma confluent; (85: axial paren- regeneration in exploited forest is much less. chyma bands more than three cells wide); (86: However, Symphonio globu!ifero shows ade- axial parenchyma in narrow bands or lines up quate regeneration under the pioneer species to three cells wide); 92: four (3-4) cells per pa- that appear after exploitation. Tenchyma strand; 93: eight (5-8) ceUs per pa- Management The survival rate in plantations renchyma strand. Rays: 97: ray width I-3 cells; may be over 80%, but early growth is slow. In a 104: all ray cells procumbent; (106: body ray forest improvement trial at 2000 in altitude in cells procumbent with one row of upright Rwanda surviving Symphonio globu!ifero and/or square marginal cells); 1/5: 4-12 rays showed a diameter growth of 0.8 cm/year and per min. Mineral inclusions: 136: prismatic length growth of 0.7 in/year and good hole form crystals present; 141: prismatic crystals In non- 32 years after planting. chambered axial parenchyma cells; (142: pris- Diseases and pests Freshly cutlogs are very matic crystals in chambered axial parenchyma susceptible to bark beetles (Plotypodinoe and cells). SCOlytinoe) and need to be processed quickly co. Ebanyenle, A. A. Oteng-Amoako & P. Baas) Harvesting Logs are felled when they are Growth and development Multiplication 50-80 cm in diameter. The exuded gum ls usu- of Symphonio globu!ifero in open locations Is ally collected from the base of the tree in the by root suckers, in forest by seed. Seedlings are form of lumps. shade tolerant, but at later stages more lightis Yield Little information on yield is available needed. Growth follows Massart's architectural from Africa. In western Gabon only 0.5 mama modelwith an orthotropic trunk with rhythmic of Symphonio g!o6ulifero timber has been regrowth and branching; the PIagiotropic corded, but in Guyana the forest contaLns on branches also grow rhythmicalIy. Early growth average 2-3 matha of harvestable timber (above is slow, about 15 cm/year. Branching starts 40 cm log diameter) when the stem is 30-40 cm long. Individual Handling after harvest Freshly harvested trees may flower for almost 2 months. They are logs may sink in water and then cannot be very conspicuous in flower. Pollination is ef- transported by river. However, the density of fected by birds (in Africa by sun birds) and freshly cut logs in Gabon is reportedIy 800-950 various insects such as wasps, bees and butter- kg/in3 which implies that they float in water. flies. Fruiting is most abundant during the dry Genetic resources Symphonioglobulifero is season The seeds are dispersed by small widespread and IOCaUy common wlth good re- mammals, e. g. monkeys, which relish the generation, and does not seem to be in danger 532 TIMBERSl

of genetic erosion. Meudon side hardness 5.7. The wood saws Prospects Symphonto globulifero is poorly slowly, with occasional charring and a strong known as a timber in tropical Africa. It may dulling effect on sawteeth; it is recommended become more important as a market already that logs be quartersawn to reduce warping. exists. It may be useful for planting to protect The wood can be finished to a smooth surface watershed areas, swamps and river banks. The gluing properties are moderate. The wood Major references Bamps, Robson & Verd- is durable, but occasionally attacked by ter- court, 1978; Burki11, 1994; CAB International, mites. The heartwood is very resistant to jin- 2005; CTRAD-Foret, 1999a; CIRAD Forestry pregnation with preservatives. Both the leaves Department, 2003; de Saint-Aubin, 1963; For- and bark contain tannins and sterols. est Product Laboratory, 1999; Katende, Birnie Botany Evergreen medium-sized tree up to & Tengn6. s, 1995; Richter & Danwitz, 2000. 30 in tall; hole often fluted at base and sinuous, Other references Abbiw, 1990; indrich & up to 170(-200) cm in diameter, often with Hamrick, 1998; Barnps, 1970; Bayma, furuda broad buttresses at base; bark surface shal- & Net0, 1998; Bras & Maury-Lechon, 1986; lowly fissured and finely scaly, inner bark or- Chapoulet & Perrier, 2001; Croptier & Kucera, ange brown to pinkish, soft-fibrous, exuding 1990; Dick, Kobinah & Bermingham, 2003; latex; young branches initially shortly hairy, Fournier, 2002; InsideWood, undated; Kabera, but soon glabrous. Leaves arranged spiralIy, 1987; Keay, 1954a; Lovett, Ruffo & Gereau, clustered at ends of branches, simple and en- 2003; Nkengfack at a1. , 2002a; Nkengfack at tire; stipules small, early caducous; petiole c. I a1. , 2002b; Robson, 1961; Takahashi, 1978; cm long; blade elliptical to obovate, 6-13 cm x White & Abernathy, 1997; Wilks & TSSemb6, 2.5-4 cm, curieate at base, rounded to shortly 2000. acuminate at apex, glabrous, pinnately veined Sources of illustration Bamps, Robson & with c. 10 pairs of indistinct lateral veins Verdcourt, 1978; Wilks & Issemb6, 2000 Flowers in fascicles on branches below the Authors L. P. A. Oyen leaves. Flowers bisexual, regular, 5-merous, small, with short pedicels; sepals fused at base, broadly ovate, shortly reddish hairy; corolla c. SYNSEPALUMAFZELii(Engl. ) T. D. Penn 35 mm long, with short tube and longer, oblong lobes; stamens inserted at apex of corolla Protologue Gen. Sapotaceae: 248 (1991). tube opposite the corolla lobes, alternating Family Sapotaceae with short staininodes toothed at margins; Chromosome number2n=28 ovary superior, globose, hairy, 5-celled, gradu- Synonyms Pochystelo microntho (Achev. ) ally narrowing into the cylindrical style. Fruit Hutch. & Dalzie1 (1931), Sersolisio microntho an ellipsoid berry c. 2.5 cm long, red when ripe, (A. Chev. ) Aub"by. & Fellegr. (1935), Afroserso- I'Seeded. Seed ellipsoid, c. 2 cm long, brown, listo gleelii(Engl. ) Achev. (1943). with very large scar. Seedling with epigeal Origin and geographic distribution Synse- germination; hypocoty12-3 cm long, epicoty1 5- polum of^elii occurs from Sierra Leone east to 6 cm long; cotyledons fleshy, c. 2 cm x ICm Cameroon and Gabon. Synsepolum o12elii flowers in C6te d'TVoire Uses In Ghana the wood is used for poles. It from December to May and fruits from June to is suitable for heavy construction, heavy floor- November Ing, mine props, vehicle bodies, sporting goods, Synsepo!urn comprises about 20 species and is railway sleepers and Implements. It can be confined to tropical Africa. It is most closely made into good-quality charcoal. The sweetly related to Errg!erophytum, which might be bet- sour fruit pulp is edible ter included in Synsepolum. The greyish wood Properties The heartwood is reddish brown, of Synsepolum stjpulotum 01adlk. ) Engl. , a the sapwood reddish yellow. The grain is usu- smalltree occurring from southern Nigeria to ally straight, texture fine. The wood is heavy, the Central African Republic, Gabon and DR with a density of about 990 kg/ms at 12% ino1s- Congo, is used for planks and axe handles. It ture content, and hard. It is difficult to dry, can also be made into good-quality charcoal with high shrinkage rates. Once dry, the wood The sweetish fruit pulp is edible and used in Is stable. At 12% moisture contentthe modulus Jams and jellies. A water infusion from the of rupture is 238 N/mm2, modulus of elasticity bark is applied as eardrops to treat earache 20,100 N/min2, compression parallel to grain 83 and a bark decoction is drunk as a galac- N/mm2, cleavage 38 N/min and Chalais- tagogue. In Nigeria Synsepolum sttpulotum

. SYNSEPALUM 533 and Synsepolum o12eliihave the same name in tagogue. Synsepolum breuipes Is occasionally some local languages. planted as a roadside tree or ornamental shade Ecology Synsepolum gleeliioccurs in lowland tree. evergreen forest, most commonly in humid Properties The heartwood is reddish brown, localities, where it is often found in small the sapwood pale brown to reddish yellow. The groups wood is heavy, with a density of about 960 Management Natural regeneration Is often kg/in3 at 12% moisture content, and hard. It abundant, with many seedlings below parent planes easily and smoothly. It splits easily and trees. The seedlings are shade tolerant. The does not take nails. It is not well suited for bole of Synsepolum o12elii may split during or turning. The wood is durable. soon after felling Botany Evergreen medium-sized tree up to Genetic resources and breeding Synsepo- 25 in tall, but usually smaller; hole straight, Ium o12eliiis fairly widely distributed and 10- strongly fluted, up to 100 cm in diameter, cally fairly common, and there are no indica- slightly buttressed at base; bark surface brown tions that it is threatened by genetic erosion to grey, scaly, inner bark pink-red, exuding Prospects There is very little information on latex; crown with spreading branches, terminal many aspects of Synsepolum o12elii, and parts drooping; young branches shortly brown evaluation of its potential as a timber tree in hairy. Leaves arranged spiralIy, clustered at managed natural forest is recommended. ends of branches, simple and entire; stipules Major references Aubr6ville, 1959d; Au- fillform, 0.5-2 cm long, persistent; petiole up to br6ville, 1964; Bolza & Keating, 1972; Burkill, I cm long; blade obovate to oblanceolate, 8-25 2000; Takahashi, 1978. cm x 4-9 cm, curieate at base, rounded to Other references Aubr6vine, 1961; Bou- shortly acuminate at apex, leathery, Initially quet & Debray, 1974; de in Mensbruge, 1966; silvery hairy below but glabrescent, pinnately Hawthorne, 1995; Keay, 1989; Kunke1, 1965; veined with 5-11 pairs of distinct lateral veins. Pennington, 1991; Voorhoeve, 1979. Flowers in fascicles on branches below the Authors R. H. M. J. Lemmens leaves. Flowers bisexual, regular, 5-meTous, greenish to creamy white, fragrant, with short pedicels; sepals free, ovate, appressed hairy; SYNSEPALUMBREVIPES (Baker) T. D. Penn. corolla 5-6 min long, with short tube and longer, oblong lobes; stamens inserted at apex Protologue Gen. Sapotaceae: 248 (1991) of corolla tube opposite the corona lobes, sta- Family Sapotaceae minodes minute or absent; ovary superior, glo- Chromosome number2n = 26, 28 hose, hairy, 5-celled, gradually narrowing Into Synonyms Foehyste!0 67euipes (Baker) Engl. the long cylindrical style. Fruit an erripsoid or (1904). ovoid berry 2-2.5 cm long, yellow to orange Vernacular names Musaka (Po). Msamvia, when ripe, with persistent style on top, I- inchocho jike, inchocha inke (Sw). seeded. Seed ellipsoid, c. 2 cm long, brown, Origin and geographic distribution Synse- with very large scar. Seedling with epigeal polum breuipes is widespread from Senegal germination; hypocoty1 0.5-2.5 cm long, epi- east to Kenya and south to Angola, Zimbabwe coty1 7-11 cm long; cotyledons elliptical, plano- andMozambique convex, I-1.5 cm long, fleshy; first two leaves Uses The wood is used for poles, stakes, opposite, with linear stipules up to 3 min long. seats, mortars and pestles, tool handles and Synsepolum 67euipes flowers in C6te d'Ivoire canoes. It is also used as firewood and for char- from September to February and fruits from coal production. The sweetly sour fruit pulp Is December to March. edible. It is also used with sugar to prepare Synsepulum comprises about 20 species and is fruit juice. In traditional medicine a root decoc- confined to tropical Africa. It is most closely tion is taken to treat malaria and as an aphro- related to Engierophytum, which inLght be bet- disiac. Sap from the roots and bark is drunk to ter included in Synsepolum. Synsepulum pobe- treat coughs, colds, hernia and stomach coin- gumionum (Pierre ex Lecomte) rug6 Assi & plaints. The leaves are used against hookworm L. Gaut. from gallery forest in West Africa is infection of the smallintestine. A bark decoc- closely related to or possibly conspecific with tion is drunk to treat swellings. The fruit pulp Synsepolum byeuipes. It is used for similar is used to treat jaundice and nausea, and the purposes. latex from the fruit is applied as a galac- Ecology Synsepolum 67euipes occurs in forest 534 TIMBERSl

as well as savanna regions, often in humid window frames and beehives. Its durability in localities along rivers and lakes, in East Africa water makes it especially suitable for boat up to 1500 in altitude. It prefers deep, well- building, and in South Africa the logs are tra- drained sandy loams rich in organic matter, ditionally used to make the jetties and slip- but is often found in localities with a high per- ways around the swamps in the Kosi Bay area inanent water table The wood is also suitable for flooring, interior Management Synsepolum 67euipes is only trim, joinery, toys, novelties, turnery, railway occasionally planted. The 1000-seed weight is sleepers, mine props, veneer and plywood. It is 1.1-1.4 kg. Seeds can be stored for some time popular as fuelwood and used for charcoal in sealed containers in a coolplace. Before sow- making. Smoke from burning wood is used to ing, seeds should be soaked in water for 12 season milk containers. hours or scarffied. In tests in C6te d'TVoire, The fruit is edible, but it has a rather bland germination started after 8-25 days, and the taste. It is popular with children. The fruit is germination percentage was high. Wildlings also made into jellies and alcoholic drinks. It are occasionally collected from the forest for yields a purple dye, whereas an orange or red- planting. The tree can be managed by coppic- dish brown dye is obtained from the bark. The ing and pollarding. In Tanzania ripe fruits are tree is a host plant of the emperor moth (Mt- collected from wild trees from October to Feb- erugone curio), of which the edible caterpillars ruary are collected. The flowers provide nectar for Genetic resources and breeding Synse- honey bees polum breujpes occurs widespread and is coin- In traditional medicine, a root decoction is mon in many regions, which means it is not drunk against amenorrhoea. Ash of the burnt easily liable to genetic erosion wood is rubbed on the forehead against head- Prospects It is unlikely that Synsepolum ache. Decoctions of the root bark and stem bark 67euipes will gain importance as a timber tree are taken for treatment of malaria. Root and because it is often too small and has a fluted bark Infusions are taken to treat cough, root hole. However, it is of interest as a multipur- and bark decoctions to treat indigestion. The pose tree providing not only wood but also bark is used as an emetic and to treat diar- good-quality charcoal, edible fruits and tradi- rhoea, stomach problems, headache, amenor- tional medicines, and it can also serve as an rhoea, wounds and respiratory problems. A leaf ornamental tree extractis drunk against cough, and an infusion Major references Aubr6ville, 1964; Burkill, of the leaves against diarrhoea and stomach 2000; Katende, Birnie & Tengnas, 1995; complaints, and as a purgative. Ground leaves, Kupicha, 1983; Ruffo, Birnie & Tengnas, 2002. bark and roots steeped in water are applied as Other references Arbonnier, 2000; Beentje, a poultice as a galactagogue. The pulverized 1994; Chhabra, Mahunnah & Mshiu, 1993; bark is sprinkled on water as fish poison Coates Palgrave, 1983; de in Mensbruge, 1966; Properties The heartwood is red-brown or Lovett at a1. , 2006; Neuwinger, 2000; Penning- pink-brown to greyish, and not clearly demar- ton, 1991; Raponda-Walker & Sillans, 1961; cated from the sapwood. The grain is usually Williamson, 1955 straight, sometimes wavy or interlocked, tex- Authors R. H. M. J. Lemmens ture fine and even The wood Is moderately heavy, with a density of 610-830 kg/ina at 12% moisture content. mr- SYZYGIUM CORDATUM Hochst. ex C. lirauss drying Is slow, with a marked tendency to dis- tort and split. The rates of shrinkage are high, Protologue Flora 27: 425 (1844) from green to oven dry 4.0% radial and 10.1% Family Myrtaceae tangential. Water-seasoning makes the wood Vernacular names Water berry, water wood, more durable. The wood is moderately hard water tree (En). Timuncho (Po). Mzambarau and moderately strong. At 12% moisture con- 21wa, myamayu, inIati(Sw) tent, the modulus of rupture is about 75 Origin and geographic distribution Syzy- N/min2, modulus of elasticity 7250 N/min2 gium cordotum Is distributed from DR Congo compression parallel to grain 45 N/mm2 and eastward to Kenya and southward to South shear 14 N/min2 Africa The wood saws easily and works well with Uses The wood is used for mortars, utensils, standard tools. Pre-boring is necessary for nail- construction, beams, rafters, poles, furniture, ing; nail-holding properties are good. Moulding

. SYZYGIUM 535 properties are good, and the wood planes to a Within Syzygium cordotum 2 subspecies are smooth surface, taking a nice polish. It glues distinguished: satisfactorily. The wood is recorded to be dura- - subsp. cordotum: shrub or small to medium- ble, particularly in water. In Zimbabwe It has sized tree up to 20 in tall, young stems dis- been recorded to be resistant to termites. The tinctly 4-angled or slightly winged, leaves sapwood is susceptible to Lyetus borers. The distinctly cordate and amplexicaul at base; heartwood is very resistant to impregnation distributed from DR Congo eastward to with preservatives, the sapwood moderately Kenya and southward to ingola and South permeable. The wood is resistant to fire. Africa, up to 2400 in altitude; Methanol and water extracts of the bark have - subsp. shimboense Verdc. : smalltree up to 7 shown in-vitro antifungal activity agalnst in tall, young stems mostly only scarcely Coildid0 o16icons. Leaf extracts have shown quadrangular, leaves slightly cordate to in-vivo hypoglycaemic activity In rats. The rounded or broadly curieate at base; distrib- bark and wood contain proanthocyanidins, uted in Kenya and Tanzania pentacyclic triterpenoids, steroidal triterpe- Syzygium cordotum hybridizes with Syzygium DoIds, erragic acid and gallic acid. The anti- gumee, !se myilld. ) DC. , and the 2 species are diarrhoealproperties of the bark may be due to connected by a complete range of intermedi- the presence of phenoliccompounds ates. Botany Evergreen shrub or small to me- Ecology Syzygium cordotttm usuaUy occurs dium-sized tree up to 20 in tall; hole up to 60 in woodland and forest, nearly always near Cm In diameter, seldom straight, often water or along watercourses, sometimes doinI- branched and gnarled, occasionally buttressed; nant in swamp forest, from sea-levelup to 2400 bark surface rough, flaking or fissured, pale or in altitude. The average annual rainfallin the dark brown; crown dense, spreading; area of distribution is usually 750-1200 min. branchlets 4-angled or slightly winged. Leaves The tree is resistant to fire. opposite, clustered near the ends of branches, Management Seeds can be used for propa- simple and entire; petiole up to 2.5(-5) mm gation. The 1000-seed weightis 2-2.5 kg. Seeds long; blade oblong, oblong-elliptical, lanceolate- are recorded to retain viahinty for a day only, elliptical or almost round, 2.5-13.5 cm x 2-8 and should not be dried in the sun. Germina- cm, base cordate and amplexicaul, rounded or tion of fresh seed is good and rapid (90% in 25 broadly curieate, apex rounded to acute or days), and pretreatment is generally not con- rarely short-acuminate or notched, blue-green sidered necessary. In an experiment in Malawi, above, paler green below, leathery. Inflores- however, germination was 93% after soaking at cence a terminalcyme up to 10 cm in diameter, room temperature for 24 hours, whereas vari- many-flowered. Flowers bisexual, regular, usu- ous other treatments gave low or no germina- ally 4-merous, white, pinkish or yellowish, fra- tion. Seeds can be sown directly In the field or grant; pedice1 I-3 min long; calyx 6-9 mm sown in pots in a nursery. Wildlings are some- long, with short lobes; petals united into a cap- times planted out. POUarding is possible. Stem like covering up to 3.5 mm x 6 mm; stamens and branch cankers have been observed, an numerous, 10-15 mm long, cream or white, caused by Chrysoporthe oustroofricono, consplcuous, fluffy; ovary inferior, 2-celled, important pathogen of EUCo!JPtus spp. world- style (5-)10-15 mm long. Fruit an oblong to wide. Several Botryosphoerioceoe fungi cause nearly globose or urn-shaped berry I-2 cm x cankers in Syzygium spp. as well as in EUCo- co. 5-)I cm, purple when ripe, tipped by the Iyptus spp. Logs often split during felling, and persistent calyx, usually I-seeded. heart rot may be present. Syzygium cordotum grows fast. In experiments Genetic resources and breeding Syzy- in Malawi, trees were 2.7 in tall 27 months ginm cordotum has a large distribution area after planting. In southern Africa flowering Ls and there are no indications that it is threat- in August-November, and fruiting in Novem- ened by genetic erosion. It is protected in South her-March Africa Syzygium is a large genus of about 1000 spe- Prospects The wood of Syzygium cordotum cies, confined to the Old World tropics and sub- is strong, easy to work and very durable in tropics, with the greatest diversity in South- water, making it particularly suitable for spe- East Asia. In the past it was lncluded in cific purposes such as boats and jetties. Be- EUgenio, which now mainly coinprlses specles cause the hole is seldom of good shape, the from the New World. commercial prospects are limited, unless

. 536 TIMBERSl

straight-stemmed types can be selected. sleepers. It is also suitable for vehicle bodies, Major references Bolza & Keating, 1972; Interior trim, Joinery, toys, novelties, boxes, Katende, Birnie & Tengnas, 1995; van Wyk & crates, mine props, veneer, plywood, hardboard van Wyk, 1997; Verdcourt, 2001; Zambia For- and particle board. It is good fuelwood and est Department, 1979d used for charcoal making. Other references Chilufya & Tengnas, The fruit is edible. It has been described as 1996; Maghembe at a1. , 1994; Musabayane at sweet and juicy, but also as having a rather a1. , 2005; Nakabonge at a1. , 2006; Neuwinger, bland taste and not being appreciated. It is 2000; Ssegawa & Kasenene, 2007; Steenkamp, sought after especially by children, and in Fernandes & van Rensburg, 2007; van Vuuren, Ethiopia and Kenya the fruit is sold on mar- Banks & Stohr, 1978; van Wyk & Gencke, kets. The fruitis made into a beverage, vinegar 2000; van Wyk, van Oudtshoorn & Gencke, and added to spirits for flavouring. In Sudan a 1997 mealis made from roasted and ground fruits. Authors M. Brink The bark is used for tanning and dyeing. Bark extracts are sometimes used to harden latentic floors or to glaze pottery. The leaves and fruits SYZYGiUMGUINEENSE myilld. ) DC. are used as fodder for livestock, and the flowers are a source of nectar for honey bees. The tree Protologue Prodr. 3: 259 (1828). is used as a shade tree in coffee cultivation in Family Myrtaceae Ethiopia Synonyms Memecylon lopezionum Achev Syzygium gumeense finds wide application in (1917). African traditional medicine, but can be dan- Vernacular names Water berry, water pear, gerous, as the bark may be poisonous, and snake bean tree (En). hisa d'eau (Fr). Mzuari, death after its use has been recorded. The root inzambarau, inzambarau inwitu, Iubale, is soaked in water for drinking and bathing to inkongoro, inlungiro (Sw). treat epilepsy. A root infusion is drunk for Origin and geographic distribution Syzy- treatment of stomach-ache. Root extracts are glum guineense Is one of the most widely dis- taken as a purgative, arithelmintic and taenia- tributed African trees, occurring from Senegal cide. Bark decoctions are used against stom- eastward to Somalia and southward to Na- ach-ache, diarrhoea and malaria; they are con- inIbia, Botswana and South Africa. It also DC- sidered mildly laxative, and are applied in curs in SaudiArabia and Yemen. draught or in baths as a tonic. hamfusion is Uses The wood is used for construction, floor- taken against coughs, asthma, throat problems Ing, panelling, furniture, utensils, tool handles, and Intercostal pain. The powdered bark is plates, stools, carvings and poles. Its flexibility used as an antispasmodic, purgative and makes it suitable for bows and ribs of canoes. arithelmintic, and used for treatment of diar- The hole is made into dugout canoes. In East rhoea, stomach-ache, broken bones and Africa the wood has been used for railway wounds. In Cameroon the bark is used for the treatment of snakebites. Twig bark preparations are applied against paralysis. A decoction

a of twigs and leaves Is drunk or used as an enema for its purgative properties and against

" A. colic, diarrhoea and abdominal pain. It is also used as drink or bath against insanity, amen- \ ,, g?^I orrhoea and cerebral malaria. The crushed leaf Is applied on wounds and boils, and taken to ILL. , a, . , treat insanity and possession. Leaf decoctions , ,A are taken against Intestinal parasites and

L. stomach-ache, used as an enema against diar- , rhoea, and used as an embrocation to bathe

** .. and then massage into areas of sprain. Leaf aa P decoctions or pulverized leaves are given as o tonic to pregnant women. The leaf is chewed against stomach-ache. A liquid of chewed leaves mixed with water Is used as eye drops to Syzygium gu"iteense - wild treat ophthalmia. The fruit is used for treating

. SYZYGIUM 537 dysentery aesthetized rats. Production and international trade No Description Evergreen shrub or small to statistics are available on the production and medium-sized tree up to BOG40) in tall; bole trade of Syzygium gumeense. The tree is of branchless for up to 15 in, seldom of good subsistence value in most parts of tropical at- shape, up to 150(-200) cm in diameter, some- rica, the timber being traded locally, mainly for times with buttresses up to 1.8 in tall; bark furniture production. The fruits are sold 10- surface smooth or with small rectangular cally flakes, grey or dark brown to almost black, Properties The heartwood is greyish red, inner bark pale brown to dark red-brown, brown or pink; it is not clearly demarcated sometimes with pink tinge or streaks; crown from the 3-4 cm wide sapwood. The grain is rounded and heavy; branchlets terete or 4- straight, texture fine to medium. Growth rings angled, sometimes drooping. Leaves opposite, are distinct. The wood is moderately heavy, simple and entire; petiole up to 2.5(-4.5) cm with a density of 640-860 kg/ms at 12% ino1s- long; blade elliptical to oblong-elliptical or ture content. Drying is slow, with moderate to obovate-elliptical, (2.5-)4-16(-17.5) cm x I-8 severe distortion and splitting. In Tanzania cm, curieate at base, obtuse to acuminate at boards of 2.5 cm thick air dry in 4 months and apex, leathery, shiny dark green above, pale boards of 5 cm thick in 7 months. Kiln drying green below, pinnately veined with many Iat- should be carried out at low temperatures; It eral veins, fragrant when crushed. Innores- gives a pronounced risk of distortion, and cence a termlnal cyme 5-19 cm long, many- checking and splitting are fairly common. The flowered. Flowers bisexual, regular, 4-meTous, rates of shrinkage from green to oven dry are white, fragrant, sessile; calyx with slender tube high: 3.6-5.9% radial and 7.8-11.5% tangen- and small teeth, persistent; petals 2-3 min tial. Movement in servlce can be large. long; stamens numerous, 4-8 mm long, white, The wood is strong. At 12% moisture content, showy; ovary inferior, 2-celled, style about as the modulus of rupture is 1/9-173 N/min2, long as stamens. Fruit a globose to ellipsoid compression paraUel to grain 43-65 N/mm2, berry 0.5-3.5 cm x 0.6-2.5 cm, red to purplish shear 8 N/min2, cleavage 16-20 N/min and black, tipped by the persistent calyx, usually I- Chalais-Meudon side hardness 30-4.7. The wood saws easily, but sawdust tends to adhere to the blades when the wood is fresh. It works easily with machine and hand tools, but is 11- able to splitting. It planes easily. Pre-boring is , necessary for nailing; nail-holding properties are good. Moulding properties are good, and the wood bores and mortises well, when done \ with care. Reports on the durability of the wood vary, but it is frequently recorded as non- II I a_. durable. The heartwood is extremely resistant --....^ I to impregnation with preservatives, the sap- \\\ I 11 .,^ wood is permeable \ Fruits conected in Malawicontained per 100 g 'L dry matter: energy 1096 kJ (262 kcal), crude protein 10.1 g, fat 4.0 g, carbohydrate 48.5 g, ., fibre 30.3 g, Ca 23 ing, Mg 225 ing, P 30 ing, Fe ,. 76 ing. They also contain ascorbic acid. 11 Extracts of the stem bark, leaves and seeds 3 have shown antibacterial and antifungal activity, and methanol extracts of the stem bark molluscicidal activity. Triterpenes with antibacterial activity against ESCherichio coli, Bo- tillus subtilis and Shigello sonnet were isolated 2 from a methanolleaf extract, the most active being an unohc acid and asiatic acid. A metha- Syzygium gumee, Ise - I, flowering twig, 2, flower, . nolextract of the bark inhibited contractions of 3, fruits. rabbit ileum and produced hypotension in an- Source: F10re attolytique du B6nin 538 TIMBERSl

seeded. Seed rounded, yellowish to brownish a tree up to 20 in tall, with a bole diameter up Other botanical information Syzygium is to 50 cm. The wood is used for panels. a large genus of about 1000 species, confined to Syzygium inichiethwoitii Verdc. (synonym: the Old World tropics and subtropics, with the Syzygium seierophyllum Brenan) is a shrub or greatest diversity in South-East Asia. In the tree up to 30 in tall, restricted to Kenya and past it was included in EUgenio, which now Tanzania. The wood is used for mortars, tool mainly comprises species from the New World. handles, utensils and poles. The fruit is edible, Syzygium gumeense is an extremely variable and the tree also supplies fodder, shade and species with a complex taxonomy. It occurs in a amenity wide range of vegetation types and shows a Anatomy Wood-anatointcal description orAWA large variety of growth forms, ranging from a hardwood codes): lofty forest tree with large plank buttresses to Growth rings: (I: growth ring boundaries dis- a rhizomatous undershrub. The flowers, how- tinct); (2: growth ring boundaries indistinct or ever, are rather uniform and the fruits show absent). Vessels: 5: wood diffuse-porous; (12: only sright difference in shape. Variation in solitary vessel outline angular); 13: simple per- leafshape and size seems to be continuous, and foration plates; 22: intervessel pits alternate; much of the variation appears to be closely 23: shape of alternate pits polygonal; 26: in- correlated with ecology and habit. The taxo- tervessel pits medium (7-10 grn); 29: vestured nomic situation Is further complicated by by- pits; 31: vessel-ray pits with much reduced bridization with Syzygtum cordotum Hochst borders to apparently simple: pits rounded or ex C. Krauss and backcrossing with the par- angular; 32: vessel-ray pits with much reduced ents. Primarily on the basis of the leaf shape, borders to apparently simple: pits horizontal 11 subspecies have been recognized for Africa (scalariform, gash-like) to vertical mallsade); as a whole, the best-known being 41: mean tangential diameter of vessellumina - subsp. gumeense: tree up to 20 in tall, leaf 50-100 pin; 42: mean tangential diameter of apex obtuse to shortly acuminate, fruit up to vessellumina 100-200 prn; 47: 5-20 vessels ICm long; from Senegal eastward to Somalia per square minimetre; (48: 20-40 vessels per and southward to Namibia, Botswana and square minimetre); 56: tyloses common. Tra- South Africa, usually occurring in woodland; cheids and fibres: 61: fibres with simple to inI- - subsp. ofromontonum F. White: tree up to 30 nutely bordered pits; 66: non-septate fibres in tall, leaf apex distinctly acuminate, fruit present; 69: fibres thin- to thick-walled. Axial up to 2.5 cm long; from Sudan and Ethiopia parenchyma: 76: axial parenchyma diffuse; 77: through East Africa and eastern DR Congo axial parenchyma diffuse-in-aggregates; 80 to Angola, Zambia, Malawi, Zimbabwe and axial parenchyma anform; 82: axial paren- Mozambique, usually growing in forest; chyma winged-allform; 83: axial parenchyma - subsp. hutllense (Hiern) Finiite: subshrub confluent; 93: eight (5-8) cells per parenchyma up to 60 cm tall, leaf apex acute to rounded strand. Rays: 97: ray width I-3 cells; (102: ray or notched, fruit up to 3 cm long; DR Congo, height > I min); 108: body ray cells procumbent Tanzania, Angola, Zambia, Zimbabwe, occur- with over 4 rows of upright and/or square mar- ring in grassland; ginalcells; 109: rays with procumbent, square - subsp. mercyocorpum (Engl. ) F. White: shrub and upright cells mixed throughout the ray; or tree up to 12 in tall, leaves relatively large 1/5: 4-12 rays per min and broad, apex short-acuminate, fruit up to co. Louppe, P. Derienne & E. A. Wheeler) 4 cm long; from Gambia to Cameroon, Ga- Growth and development Pollination is by bon, Central African Republic and DR Congo, insects. Natural regeneration is adequate in in woodland and seasonalIy wet grassland. natural forest The wood of various other Syzygium species is Ecology Syzygium guLneense occurs up to used. Syzygtum 60rbonicum J. Gu6ho & 2700 in altitude, in areas with an average tern- A. J. Scott is endemic to Reunion, where it is perature of 10-30'C and an average annual known as 'bois de pornme', 'bois de pornme rainfall of 700-2300 min. It is found in a wide blanc','hots de p6che marron' or 'bois a 6corce range of vegetation types, including rainforest, blanche'. It is a buttressed tree up to 20 in tall. montane forest, riverme forest and woodland The wood is used in construction It prefers moist, well-drained soils with a high Sy2ygium cymosum (Lain. ) DC. is endemic to water table. The presence of the tree is consid- Reunion and Mauritius, where it is known as ered a fairly good indication that the ground- 'bois de pornme' or 'bois de pornme rouge'. It is water table is near the surface

. TABERNAEMONTANA 539

Propagation and planting Propagation by Tesemma, BITnie & Tengi6. s, 1993; Coates seed is easy and is the commonly used method Palgrave, 1983; Djoukeng at a1. , 2005; HUSsaln of propagation. The 1000-seed weight is 270- & Deeni, 1991; InsideWood, undated; Katende, 420 g. The seeds must be sown Jinmediately Birnie & Tengnas, 1995; Lovett at a1. , 2007; after the fruits are picked, as they may spoil Malele at a1. , 1997; Neuwinger, 2000; Oketch- within 24 hours of storage. Germination is Rabah & DOSsaji, 1998; Saka & Msonthi, 1994; usually very good and uniform, and pretreat- Scott, 1993; Sommerlatte & Sommerlatte, merit of the seed is not necessary. Germination 1990; Ssegawa & Kasenene, 2007; Tanzania takes 20-30 days, with a germination rate of Forest Division, 1963b; Tchi6gang-Megueni at 80-90%. Seeds can be sown directly in the field a1. , 2001; van Vuuren, Banks & Stohr, 1978; or are sown in pots in a nursery. Wildlings can van Wyk & van Wyk, 1997; White, 1977; Wim- also be used for planting. Stem cuttings root bush, 1957. easily, and grafting has been tested with a suc- Sources of illustration Ako6gninou, van cess rate of 50% der Burg & van derMaesen, 2006 Management The tree tolerates pollarding Authors A. Maroyi and is able to coppice. Diseases and pests Syzygium gumeense Is liable to attack by larvae of a Cerambycid bee- TABERNAEMONTANASTAPFIANABritten tle, which can make the timber defective. Heart rot is often present in larger trees. Stern Protologue Trans. Linn. SOC. London, Bot. 4 and branch cankers have been observed, 25 (1894). caused by Chrysoporthe oustroofricono, an Family Apocynaceae important pathogen of EUCo!yptus spp. world- Synonyms Tobernoemontono johnston, , wlde (Stapfj Pichon (1948) Harvesting Ripe fruits are generally picked Vernacular names Wild magiioha, soccerball from the tree. If fallen, they must be picked fruit (En). Mwambe (Sw). immediately from the ground so that they do Origin and geographic distribution rober- not spoil ridemontono stopfiono occurs from eastern DR Genetic resources Since Syzygium gumeense Congo to eastern Kenya, and south to Zim- is widespread and rather common, it does not babwe and Mozambique. seem to be threatened. In countries where it is Uses In eastern DR Congo the wood is used not common, such as Ghana, it may deserve for construction of local houses, combs, spoons some protection. and handles for knives and swords. It is also Prospects Syzygium gumeense may have used as firewood. The bark, seeds and roots are some prospects as a timber tree for local use in used against high blood pressure. The latex Is tropical Africa, as it is widely available and the applied to wounds. In Kenya a leaf decoction is wood is hard, strong, easy to work and suitable used as an appetizer and to treat colic, and in for a range of uses. Commercial prospects are DR Congo and Burundi as a galactagogue and limited, because the hole is seldom of good in strong doses as an abortifacient. The latex Is shape. The vanability within the species can be used as birdhme. Tabernoemontono stopfiono used for selection and cloning of superior geno- is planted as an ornamental, and as a shade types. More detailed knowledge of selection tree and bee plant. and standardized vegetative propagation meth- Properties The wood is soft and white to ods are the main prerequisites for a break- yellow or pale brown. through in productivity. Although the tree is About a dozen indole alkaloids have been ISO- widely used in traditional medicine, caution Is lated from the bark and stems of Tabernoe- needed as death has occurred after use of the montono stop/tung. These include perlvine, bark. Research on the phytochemistry is war- tubotaiwine and Ibogamine, and several phar- ranted. macologically active derivatives. Major references Bolza & Keating, 1972; Botany Small to medium-sized tree up to Bryce, 1967; Burki11, 1997; Eggeling & Dale, 25(-35) in tall, glabrous; hole up to 90 cm in 1951; Palmer & Pitman, 1972-1974; Takaha- diameter; bark rough, pale to dark grey-brown; shi, 1978; Verdcourt, 2001; White, 1978; Wil- crown widely spreading. Leaves opposite, SIm- namson, 1955; World Agroforestry Centre, un- PIe and entire; ocrea conspicuous, usually wid- dated ened into stipules in axils of petioles; petiole 5- Other references Beentje, 1994; Bekele- 30 min long; blade narrowly elliptical, 1240 540 TIMBERSl

cm x 3-14 cm, base curieate, often decurrent being useful not only for its timber but also in into the petiole, apex acuminate to rounded, traditional medicine and as an ornamental pinnately veined with 12-24 pairs of lateral Major references Dharani, 2002; Kingston veins. Inflorescence a rather lax corymb 10-28 at a1. , 1978; Leeuwenberg, 1991; van Beek at cm long, few- to many-flowered; peduncle 3-15 a1. , 1984; Yamada, 1999 cm long, rather robust. Flowers bisexual, regu- Other references Beentje, 1994; Hyde, Iar, 5-merous, sweet-scented; pedice1 5-30 min 2002; Lovett at a1. , 2006; Neuwinger, 2000; long; sepals almost free, orbicular to oblong, 5- Njunge, 1996; 0min0, 2002; 0mino & Kok- 7 min long, thick and fleshy; corolla tube al- war0, 1993 most cylindrical, 21-42 mm long, twisted, pale Authors R. H. M. J. Lemmens green often with pale yellow throat, widened around the arithers, inside hairy, lobes obliquely elliptical, 17-60 min x 8-35 mm, un- TECTONAGRANDIS L. f dulate, spreading, apex rounded, white; stamens inserted about halfway the corolla tube, Protologue Srippl. PI. : 151 (1782) included, arithers sessile; ovary superior, al- Family Verbenaceae (APG: Lamiaceae) most cylindrical, consisting of 2 separate car- Chromosome number2n= 36 pels connate at base, style cylindrical, 5-10 Vernacular names Teak, Indian oak (En). mm long, pistil head composed of a basal ring, Teck (F"). Teca (Po). Msaji(Sw) 5 almost orbicular lobes and a stigmoid apical Origin and geographic distribution Teak part. Fruit consisting of 2 separate subglobose originates from the tropical forests of southern follicles up to 20 cm in diameter, dark green and south-eastern Asia, between 9-26'N and with paler dots, indehiscent, several- to many- 70-100'E, where it may form almost pure seeded. Seeds obliquely erripsoid, 1.5-2 cm stands. Naturalstands are found in Myanmar, long, grooved, covered by minute warts, dark India, Thailand and Laos; also in small areas brown, anIwhitish. of China and Cambodia. Teak is one of the Tuberridemontono comprises about 110 species most commonly planted species in the tropics, and is pantropical. About 18 species occur in between 30'N and 23'S. In tropical Africa it mainlandAfrica and 15 in Madagascar. was introduced mainly by the Germans, in robernoemontonu penduli/10ro K. Sebum. is a Nigeria (1889), Tannania (1898), Ghana (1905), shrub or small tree up to 8(-12) in tall, occur- Cameroon and Togo (1907-1912). From the ring from Nigeria to DR Congo. In DR Congo plantations in Togo seed was introduced into the trunk is used for house posts, and the wood neighbouring countries such as C6te d'TVoire, is also used for small blackboards as it is so where the first large plantations were estab- soft that letters written on it can be easily lished in 1929 in the savanna zone near erased; the wood serves as firewood. The latex Bouak6, and Benin and Senegal in 1933 is used as birdlime and in traditional medicine Uses Teak has been a highly appeciated to treat colds and wounds. In Cameroon the timber for centuries in its region of origin. roots are used to treat malaria. In the Central African Republic fruit juice is applied to wounds, and the bark is made into rough rope. The main flowering period of Tuberitoemon- .. toner stopfiono is December to February Ecology Tobernoemontono stopfiono occurs @ ^ in forest at 700-2500 in altitude. It is coin- , monly found in secondary forest and on river banks. 7 .. - Management In forest in Kenya, more than 5 Tuberitoemontono stopfiono stems of over 20 @ cm diameter can be found locally per ha. Genetic resources and breeding Tuber- 00 Floemontono stopfiono is not threatened by , genetic erosion because it is common even in o disturbed forest. Prospects Tuberrigemontono stopfiono deserves more attention as a multipurpose tree, Teetonogrondis -pionted

. TBCTONA 541

Wooden sculptures, doors and coffins made of mand for timber, and it is the timber most in teak and over 1000 years old have been found demand. In Africa the largest exporter is C6te in good condition in Indian and Persian tern- d'Ivoire with over 100,000 ina of logs per year PIes. Under cover, teak is considered virtuaUy (127,000 ms in 1998), followed at great distance imperishable. It is the preferred wood for naval by Ghana, Nigeria, Togo (1700 ing during the construction, elements of boat hulls and first 4 months of 2004) and Cameroon (40 ina of bridges of boats because of its extreme resis- sawn wood in 2003). Large-diameter logs from tance to decay and corrosive activity of water. natural forest in the region of origin may fetch It is suitable for a variety of uses: luxury cabi- very high prices (in 1998 logs for veneer pro- net-work, frames (doors and windows), furni- duction fetched Us$ 1700-2000/in3 and sawn ture and joinery with solid wood, parquet floor- wood Us$ 600-1300/ms), which makes teak the ing, staircases, carpentry, garden furniture, luxury wood par excellence. Timber originating railway sleepers, bridges and other construc- from plantations fetches lower prices, but be- tion in contact with water or the ground, elec- tween 1996 and 1999 the price (FOB) in C6te tricity and telephone poles, fence posts, rail- d'TVoire reached Us$ 2501m3, after which it fell cars, barrels and vats for liquids, even corro- A small part of the African production is ex- SIve ones, various utensils (except tool handles ported to Europe, but the major part goes to because the wood is brittle), turnery and sculp- India, which has equipment available that is turing. Through slicing teak makes nicely fig- adapted to logs of small dimensions as well as ured veneer. It is also used as firewood. It is ample manpower. India is the largest importer often planted by farmers around fields as living of teak, but also the largest exporter of proc- fence posts and to support barbed wire. It is essed teak wood. also planted as an amenity tree in towns and Properties The heartwood is brown-yellow, along roads. darkening and developing a golden shine with The leaves are used for packing food products age. It is sometimes beige and may show in the markets. The root bark and young leaves darker veins, and occasionally is completely yield a yellowish brown, red or yellow dye, brown-black due to regular exposure to bush which is used to colour paper and textiles (silk, fires, the latter condition generally riot being cotton or WOOD, but which needs a mordant appreciated by the wood-processing industry. It The dye from the leaves prevents blackening of is distinctly demarcated from the whitish sap- yam after cooking. The leaves have a reputa- wood, which is (I-)2-6 cm wide. The grain is tion of being diuretic, depurative, stimulant, usually straight, texture rather coarse. Growth antidysenteric and vermifuge, and are used in rings distinct. The wood is oily to the touch and traditional medicine to treat anaemia, asthe- has a characteristic smell of old leather when nia, fever and malaria, amoebiasis, schisto- fresh. somiasis and tuberculosis. All oil extracted Teak is a medium-weight wood. At 12% inois- from young shoots ls said to be used as a ture content, the density is (480,610-730(- treatment for scabies. The flowers are used 850) kg/ina. The rates of shrinkage are low, against billous affections, bronchitis and un- from green to 12% moisture content 0.7-1.5% nary complaints; they are diuretic, as are the radial and 1.1-2.5% tangential, and from green seeds. The seed o111s applied as a hair tonic to oven dry 1.33.7% radial and 3.2-6.2% tan- The astringent bark is used to treat bronchitis gential. Apparently, the rates of shrinkage are The sawdust is used for the production of ac- not influenced by the growth rate of the tree tive carbon. Teak dries very well but rather slowly. Boards Production and international trade A1- of I cm thick take 15 days to air dry from 40% though statistics are rather incomplete, it is to 15% moisture content, boards of 2.5 cm thick estimated that teak plantations cover 5.7 inil- 30 days and boards of 4 cm thick 50 days lion ha worldwide. In tropical Africa, the area Boards of 2.5 cm thick can be kiln dried from under teak plantations is about 250,000 ha, the 40% to 10% moisture content in 5-6 days at a most important countries being Nigeria (70,000 drying temperature of 60-80"C, and a corre- ha), C6te d'TVoire (60,000 ha), Benin (21,000 sponding relative humidity of 80% to 40%. Par- ha), Ghana (17,000 ha), Togo (16,000 ha), Su- ticular care is required in determining initial dan (14,000 ha), Tanzania (10,000 ha) and and final moisture contents, as large variations Senegal(2500 ha). in drying rates occasionally occur. The wood is Currently, the world production of teak (about very liable to colour change, and high initial 2 million ms/year) covers I% of the world de- temperatures should be avoided. During kiln 542 TIMBERSl drying a condensate of butyric acid is formed, tion. The fine dust produced in machining op- which may corrode metal kilns (unless made of erations may cause irritation of the skin or aluminium or stainless steel). Teak has low bronchial asthma and rhinitis after inhalation; movement values and a good form stability a well-functioning dust extractor fan is recoin- At 12% moisture content, the modulus of rup- mended. The substance responsible for the ture is 81-196 N/min2, modulus of elasticity allergic reaction is probably the naphtho- 7600-17,500 N/min2 compression parallelto quinone desoxylapachol. mother naphtho- grain 34-70 N/mm2, compression perpendicu- quinone, lapachol, isolated from the roots was Iar to grain 6-8 N/mm2, shear 5-16 N/min2, found to have an anti-ulcerogenic effect on ex- cleavage 14-30 N/mm, Janka side hardness perlmentally induced gastric and duodenal 3700-4890 N and Janka end hardness 4065- ulcers in rats and guinea-pigs. Wood extracts 4760 N. showed potent leishmanicidal activity. The Teak is not difficult to work, but requires some resistance of teak wood to termites and fungiis effort, mainly because of the presence of silica due to the presence of tectoquinone and other (up to 1.5%). Stentte-tipped saws and tools arithraquinones. The energy value of the wood tipped with tungsten carbide are indispensable is 21,350 kJ/kg. The seed kernelcontains about for sawing and planing operations. The wood is 40% oil. difficult to chisel with a hollow square inor- Adulterations and substitutes The wood tiser, but turns well. The nail- and screw- of several tree species indigenous to Africa is holding capacities are good, but pre-boring is traded as 'African teak': Bothioeo PIur!jugo recommended to avoid splitting. Gluing is only Harms (also called Zarubeziteak or Rhodesian successful on freshly machined or newly teak), Miltcio excelso orelw. ) C. C. Berg (Iroko), sanded surfaces. The wood bending properties Oldfieldi0 o17icono Benth. & Hook. f. (also are moderate; there is a tendency to buckle, called African oak) and Pencopsis e!oto and the wood is only suitable for bends of inod- (Harms) Meeuwen (afrormosia). erate curvature. Painting, staining and polish- Description Deciduous medium-sized to large ing require freshly machined surfaces or a pre- tree up to 40(-50) in tan; bole generaUy straight treatment with thinner. However, teak can be and branchless for up to 20 in, up to 150(-200) varnished and polished beautifully. his easy to cm in diameter, at base fluted or with low but- cut into smooth, tight veneer of uniform thickness at a temperature of 90-95'C. The veneer 5 dries flat and split-free with low shrinkage Owing to its beautiful figure, teak is in great demand as face veneer. The wood can be t;\ 111.1 peeled, but slicing gives the most beautiful and ';) I-.',,.,!. I 4 I varied veneer patterns. 531, Teak heartwood is rated as durable to very I. \ durable. Stake tests showed an average service 2 ? life in contact with the ground of more than 10 J! years under tropical conditions and more than I ,. * b 25 years under temperate conditions. However, .* ^ ;- there are indications that the durability is SIg- , nificantly influenced by tree age. Teak is very , <;* durable under cover. The heartwood is resis- 1:9 .~*~ tant to termite and fungal attack, and resistant ,* but not immune to marine borer attack. The I sapwood is reported to be liable to attack by powder-post beetles (Lyetus sp. ). The heart- 3 J wood is difficult to treat with preservatives, , I 1.1 but treatment is usually only needed for the *" sapwood. -*3- The wood contains 38-47.5% cellulose, 30-33% 11gnin, 12.5-14.5% pentosan, 1.1-1.4% ash and Tectonogrondis - I, tree hubit, 2, flower, 3, fruit- 0.2-1.5% silica; the solubility is 4.6-10.7% in ing twig, . 4, fruits enclosed by colyx, . 5, fruit alcohol-benzene, 1.2% in cold water, 11.1% in with opened calyx. hot water and 15.6-19.8% in a I% NaOH solu- Source. PROSEA

. TBCTONA 543 tresses; bark longitudinalIy fissured and finely (N. P. Monel & P. Baas) scaly, grey to brown; crown rounded, open; Growth and development Teak is a strong young branches tetragonal, rusty brown hairy. light demander and a long-lived pomeer that Leaves decussately opposite, simple and entire; may be invasive. It is said to be sensitive to stipules absent; petiole 2.5-5 cm long, hairy; competition at the level of the crown and of the blade broadly ovate to elliptical-ovate or roots, but it can be associated with species hav- obovate, 20-60(-100) cm x 15-35(-50) cm, ing a deep root system. In deep soils teak de- curieate at base, acuminate at apex, thinly velops a taproot, which makes it resistant to leathery, glabrescent above, hairy and scabrous winds. In more shallow soils teak develops with below, pinnately veined. Inflorescence consist- age several very strong superficial roots, which ing of cymes arranged in a large terminal pani- are concentrated in the upper 50 cm of the soil cle up to 70 cm long. Flowers bisexual, regular, and may extend up to 20 in from the stem, en- 5-7-merous, small, 3-6 mm long; calyx shortly sunng its stability. Its fibrous bark makes it lobed, persistent; corolla with a short tube and resistant to fires, which are favoured by the longer, patent or reflexed lobes, white with thick layer of easily inflammable litter. Some pink on the lobes; stamens inserted at the base claim that fires promote the growth of teak of the corolla tube; ovary superior, ovoid, 4- Fires may break seed dormancy and favour celled, style slender, stigma 2-lobed. Fruit a germination. From a very young age teak globose drupe I-1.5 cm in diameter, densely sprouts abundantly after fire has passed or hairy, usuaUy I-2-seeded, enclosed by the ac- after felling, and the sprouts form undergrowth crescent calyx. Seeds ovoid, c. 6 mm x 4 mm, in thinned populations. It is possible to grow a without endosperm. Seedling with epigeal forest from tree stumps, resulting from re- germination; cotyledons with a notched apex. growth after final exploitation by clear cutting Other botanical information Teetono coin- The abundance of sprouts can be a problem prises 4 species, all from tropical Asia; the po- when replanting with new Improved planting sition of the genus within the family seems material rather Isolated Teak is a deciduous species. Its phenology de- Anatomy Wood-anatointcal description OAWA pends on the climatic zone in which it is hardwood codes): planted. The sprouting of buds during the first Growth rings: I: growth ring boundaries dis- rains precedes a period of fast apical growth tinct. Vessels: (3: wood ring-porous); (4: wood (one month). Flowering takes place after the semi-ring-porous); 13: simple perforation period of growth in height, followed two plates; 22: intervessel pits alternate; (23: shape months later by the maximum Increase in di- of alternate pits polygonal); 25: intervessel pits ameter of the trunk. In plantations, the first small(4-7 prn); 26:intervesselpits medium (7- flowering occurs from the second or third year 10 F1m); 30: vessel-ray pits with distinct bor- As the inflorescences are terminal, they fre- ders; similar to intervessel pits in size and quently cause forking of the trunk, which shape throughout the ray cell; 42: mean tan- sometimes disappears when the tree grows gential diameter of vessellumina 100-200 prn; The height under the fork, representing the (43: mean tangential diameter of vessellumina height of the exploitable trunk, depends thus Z 200 F1m); 47: 5-20 vessels per square minime- in general on the earnness of first flowering. tre; 56: tyloses common; 58: gums and other An inflorescence can have up to 1000 buds, but deposits in heartwood vessels. Tracheids and they will not allflower. The flowering of a sin- fibres: 61: fibres with simple to minutely bor- gle inflorescence is spread over 46 weeks, but dered pits; (65: septate fibres present); 66: non- each flower lasts for just one day. The amount septate fibres present; 69: fibres thin- to thick- of pollen released is highest when the stigma is walled. Axial parenchyma: 79: axial paren- receptive, around noon. Insects, especiaUy chyma vasicentric; 89: axial parenchyma in bees, take care of pollination. The level of marginal or in seemingly marginal bands; 92: autogamy is very low. In C6te d'TVoire full four (3-4) cells per parenchyma strand; 93: flowering takes place during the long rainy eight (5-8) cells per parenchyma strand. Rays season (May-August(-September)). Only a 97: ray width I-3 cells; (98: larger rays coin- smallpercentage of the flowers give fruits. The monly 4- to 10'seriate); 104: all ray cells pro- fruits attain their maximum size about 50 days cumbent; (106: body ray cells procumbent with after pollination and are mature between 1/5- one row of upright and/or square marginal 195 days after pollination. The fruiting period cells); 1/5: 4-12 rays per min. is 3-4 months(September-December) 544 TIMBERSl

In good conditions and with good provenances, 1200 in altitude. the initial growth is very fast and self-pruning Propagation and planting Because of the good. Nevertheless, as teak is sensitive to coin- spread of flowering and fruit ripening, it is petition, its growth can slow down quite rap- recommended to collect the fruits from the idly in unthinned populations, and therefore ground in order to avoid harvesting immature the first thinnings have to be early. With good seeds. The seeds show orthodox behaviour and SIIviculture in the best class of fertility, a well- can be kept at room temperature for 2-5 years. managed plantation in C6te d'Ivoire produces a For longer storage, it is advisable to keep them volume of about 560 in3ftia of wood when 35 in a cold room (4'C). The teak 'seed' is actually years old. In poor soils or in bad rainfall condi- a fruit which can contain up to four 'real' seeds. tions, growth is slow, the trunk short and the There are 800-2000(-3500) fruits per kg. Usu- branches thick, and the production can be less ally there are one or two viable seeds per fruit, than 200 mama after more than 60 years. In but some fruits are sterile. The germination of extreme conditions, teak can survive many a single seed lot may start after about 10 days years in the form of bushwood. and stop several years later; the optimum ger- Quite often teak trees have a number of defects mination rate is reached at 35-45 days. It is such as marked grooves that can reach quite quite difficult to break the seed dormancy and high on the trunk and dents that can some- so far no reliable technique has been devel- times be very abundant, strongly reducing the oped. The usual method consists of soaking the yield of sawn wood. Low forks limitthe volume seeds in water for two days, then drying them of timber. Sudden exposure of trunks to light at in the sun for a whole day, and repeating this the time of thinning may lead to the appear- for a period of 2-3 weeks. It has been shown ance of a large number of water sprouts, which that large seeds germinate better and faster damages the quality of the wood than small ones and that seed stored for longer Ecology Tropical climates are favourable for than one year shows a higher germination rate. teak if the average inlnimum temperatures do In order to stimulate germination, the seeds not fall below 18*C, if there is a marked dry should not be covered completely with soilor be season of at least two months and if annual shaded. The germination rates are approxi- rainfallis 1000-2500 mm; the longer the dry mately 20-35%. In C6te d'Ivoire, one kg of season, the better teak tolerates high rainfall. seeds generates an average of 600 plants. Teak can tolerate a marked drought in alluvial The grafting of teak, mostly shield-budding plains when groundwater is accessible for the and occasionally split-grafting, is quite easy roots. Nevertheless it is a very adaptable spe- and the rate of success is almost 100%. This cies that can survive with rainfall as low as technique is mainly used to establish clonal 500-600 mm/year or as high as 5000 mm/year. seed gardens, but the stock frequently sprouts The rainfall pattern can have one or two rainy and the 'jutcit' shoots must be removed. Layer- seasons per year. Teak can also grow in areas ing is possible, but hardly an option for the without a distinct dry season, but will then be production of large numbers of plants. Teak more susceptible to diseases such as root rot, can be propagated by cuttings and several tens which explains the bad results obtained in Ga- of thousands of plants are produced this way bon. It is frost-sensitive, but tolerates very every year in C6te d'Ivoire by Sodefor (Societe light freezing. It tolerates repeated passages of de Developpement des Forets). Teak can also fire be propagated by in vitro culture, but this Teak can grow on many kinds of soils except technique is apparently not yet used in Africa. those that are subject to prolonged or perma- Young plants originating from seeds can be nent waterlogging. Nevertheless, teak is a very grown in a seedbed (50 in' of seedbed and 6000 demanding species, which, for optimum seeds needed for one hectare plantation). Teak growth, requires a deep, well-drained soil with can easily be planted as stumps with a rate of a high mineral content, especially in Ca, Mg success exceeding 90% even without rain for and P. It prefers soils with a neutral or slightly 10-15 days after planting. Stumps can be acid pH. It develops badly on clay, on latentic stored some weeks before planting by heeling soils with low permeability and especially in them in a shady place or in moistened jute hydromorphic areas. The best results are ob- sacks. It is not really necessary to give the tamed on alluvial sandy loams or sandy loamy roots a mud bath before planting. In Asia spe- clays, that are deep and well-drained. Teak cial methods allow the stumps to be stored for plantations can be found from sea-level up to several months before planting, which makes it

. TECTONA 545 possible to prepare them before the dry season present even more optimistic internal rates of and to plant them as soon as the first rains return. begin. The planting of stumps encourages fork Although teak is tolerant of fire, fires of the formation. Plants derived from in vitro culture ground cover must be avoided to preserve the or cuttings are mostly grown in pots. The pro- long-term productivity of teak plantations, duction in pots increases the risk of inalforma- because they leave the soil bare and cause ero- tions of the root system without markedIy jin- SIOn proving the rate of success at planting. The Diseases and pests Attacks by several fungi, trees have to be planted very densely: about some actually killing the plants, have been 1500/ha, which corresponds to a 2.5 in x 2.5 in recorded. In Africa, teak plantations do not arrangement, so that they can grow straight in seem to face serious diseases apart from root search of light. For clonal plantations, a plant- rot, which may cause severe damage, especially ing distance of 4 in x 4 in (625 plants/ha) is in the south of C6te d'Ivoire and Benin, but recommended also in Nigeria and Tanzania. The fungal para- The young plants do not tolerate competition sites are Armilloriu sp. , Phellinus norius, by herbs or being shaded for too long by a can- Phoeolus monthotis, Gunodermo spp. and Ri- opy, even if the canopy is not dense. Fertilizers gidoporus Jignosus. The last one is potentially at planting enable the trees to dominate the the most dangerous. Infestation of the root weedy vegetation rapidly. During the first year system causes open places in the plantation. In 3 weedings are recommended, 2 during the Benin infestation rates vary from I% to ap- following year and a final one during the third proximateIy 60% of the trees. In Benin infesta- year. Damage to the roots during weeding tion by Botryodiplodio theo670moe causes die- should be avoided because teak is very sensi- back in patches, as does infestation by Pond tive to root rot, of which the negative effects sp. , whereas Corticium sp. kills young only become visible many years later. Teak branches. Fungal attacks causing damping-off lends itself very well to the taungya system may occur in nurseries, although teak is less and to intercropping, both of which reduce the sensitive than many other species. In zones establishment costs of plantations. Wider with high rainfall and a high water table, teak planting densities are then applied to permit suffers from a gum-disease ('water blister'), intercropping, and thinning of the lower which affects the wood quality branches is done from the second or third year In its centre of origin, more than 180 insect onwards species have been found on teak, but only some Management Some say that the growth rate of them (defoliators and borers) cause serious has little influence on the physical properties of damage. Trunk borers of the genus APGte teak wood; others state that a diameter incre- (BOStrichidoe), are reported on healthy teak merit of 1.2 cm/year gives optimal wood quality, Loro, Ithoceoe parasites have been found on while an increment of less than 0.5 cm per year teak in many countries, sometimes severely leads to inferior wood. Presently there is a ten- reducing the growth of the trees. Pests are rare dency to reduce the turn-over period to less in nurseries; defoliating or leaf-eating caterpil- than 25 years, while traditionally teak is bar- Iars are sometimes present, as well as termites vested at 35-80 years. Thinnings have to be on the roots. Rodents may constitute some done early, rigorously and frequently. In C6te danger during the germination of the seeds; d'Ivoire, a first thinning is recommended when baits or traps can be used to reduce the risk the ground surface of trees reaches 13-14 Harvesting Usually, several thinnings are mayha, and a second one when it reaches 15-16 carried out in a tree population and the final in'/ha. The tentative SIIvicultural standard Is harvest is by clear cutting. Independently of 3-6 thinnings 3-12 years apart and complete the growth rate, the turn-over period deter- felling at the age of 35-80 years depending on mines the proportion of heartwood: at 13 years soil fertility and the desired final diameter of 35-55% of the wood is heartwood, at 21 years the tree. The final felling represents 80% of the 55-65%, and at an age of over 50 years more income, wood of the firstthinnings being of low than 80%; simultaneously, the value per in310g value. In C6te d'TVoire it is estimated that the increases proportionally. In spite of the present internal return rate of a plantation under the tendency to fell the trees after about 20-25 best fertility and with a felling age of 35 years years, it seems better from a technological exceeds 10%; the return might even be 15% in point of view to stick to an average turn-over the case of clonal plantations. Other countries period (about 40 years) 546 TIMBERSl

The seeds are collected from the ground, from The genetic bentability of growth in diameter which alllitter and weedy vegetation is re- is low 01'=0.41), the bentability of the absence moved. Branches can be shaken to free the ripe of fork formation is average 01"=0.63), the fruits, but fruits are not picked from the tree to heritability of trunk straightness is high avoid the harvesting of immature seeds. 01'=0.88) and the bentability of thin branches Yield Under very good conditions in Nigeria, is very low. The heritabilities of earnness of an exceptional production of 24 matha/year has flowering and profuse fruiting are very high. In been reported. In C6te d'TVoire yields of 10-16 C6te d'Ivoire plus trees are propagated by cut- in31ha/year are recorded in the forest zone, 7- tings; they are selected on the basis of the late 10 mama/year in the transition zone and 5-7 occurrence of forking, thin branches, natural mama/year in the savanna zone. In Tanzania lopping and cylindrical hole. Up to now, breed- the best plantations are said to produce up to ing programmes do not seem to have attached 14-17 in8thatyear and in a provenance trial sufficientimportance to the wood quality yields of 12.2-19.6 mama/year were obtained at Prospects Teak is one of the best timbers the age of 17 years and existing plantations are unable to meetthe In seed gardens, the production of seeds demand. Its numerous applications, its easy amounts to 2-4(-10) kg per tree, depending on propagation and its resistance to bush fires the year make this tree more and more appreciated by Handling after harvest Teak is a heavy rural communities. Although its silviculture wood when green, and is not buoyant; it is has been mastered, the technology is not surfi- therefore transported by truck, ship or train ciently popularized to be able to optimize the Before storing the seeds, the papery remain- quantity and quality of its production. Never- ders of the calyx are removed by pounding and theless, research on the SIIviculture of clones winnowing, then the seeds are sun-dried. The must be continued because shortening the seeds are urticant and allergenic. turn-over period may reduce the timber quality Genetic resources In Asia the regression of and harm the timber's good image. Teak has a natural teak forests has been very drastic and great future ahead, but it is necessary to obtain rapid; Thailand, one of the countries of origin a good match between planting material and of teak, started importing teak wood in the plantation site early 1980s despite the establishment of the Major references Bail16res & Durand, first plantations in 1910. However, due to the 2000; Bekker, Rance & Monteuuis, 2004; Bonal fact that there are many plantations in the & Monteuuis, 1997; Dupuy & Verbaegen, 1993; world and especially thanks to the trials put in Gang10, 1999; Ganglo, Leioly & Pipar, 1999; place by FAO and Danida since 1969, coinpar- Madoffe & Maghembe, 1988; Maldonado & ing 75 provenances from 16 countries (region of Louppe, 1999; Phengklai at a1. , 1993; Rance & origin and introductions), the genetic diversity Monteuuis, 2004 of the species is not threatened. Molecular ge- Other references Bhat, 2000; Behaghel, netic analyses confirm that there are at least 1999a; Behaghe1, 1999b; Brunck, 1959; Chollet, two major groups: the very diverse Indian 1956; CTFT, 1950; CTFT, 1990a; Dupuy, 1990; group, which is subdivided into populations Dupuy, Maitre & N'Guessan Kanga, 1999; Goh from the humid, the semi-dry and the dry & Galiana, 2000; Hock & Mariaux, 1984; In- zones, and the more homogeneous Thai group, sideWood, undated; Keiding, 1985; Kjaer, including some populations planted in Indone- Lauridsen & Wellendorf, 1995; Maltre, 1983; SIa and Africa. Monteuuis, Bon & Goh, 1998a; Monteuuis, Bon Breeding Selection of a good provenance is & Goh, 1998b; Monteuuis & Goh, 1999; Mon- essential for the success of a plantation pro- teuuis, Vanauri & Chauvi6re, 1995; Souvan- gramme in a certain environment. Because of navong, 1983; Takahashi, 1978; Wericelius, the large vanability in planting material, sev- Malagiioux & Delaunay, 1975. eral countries started breeding programmes Sources of illustration Phengklai at al. , during the 1960s, which have led to the crea- 1993 tion of seed gardens in La Sangou6, C6te Authors D. Louppe d'TVoire (clones from plus trees selected from several provenances), in Gainbari, Nigeria and in IQ. roka, Tanzania. These countries possess also seed-producing populations obtained by mass selection in the finest plantations.

. THESPEsiA 547

THESPESIAPOPULNEA (L. ) Sol. ex Corr6a. caulking boats. In West Africa and India the leaves are used for wrapping food. Young Protologue inn. Mus. Natl. Hist. Nat. 9: shoots are used as fodder and as green manure 290 (1807) In South-East Asia and India the young leaves, Family Malvaceae flowers and unripe fruits are eaten raw, boiled Chromosome number 2n = 24, 26, 28 or fried as a vegetable. The flowers and fruits Vernacular names Portia tree, Pacific rose- yield a water-soluble yenowish dye, while the wood, umbrella tree, Indian tulip tree (En) wood soaked in water gives a solution that is Motel debou, feuilles d'Haiti, proch6, kalfata, used in Asia to dye wool deep brown. In Tuvalu milo, arbre ombrelle, bois de rose d'Oceanie the leaves are used for making a black dye. The (Fr). Pau rosa, bela sombra, tespesia (Po). Mta- bark has been used for tanning leather and kawa (Sw) yields a thick gum which is not soluble in wa- Origin and geographic distribution Thespe- ter. Oilfrom the seed can be used as lamp oil. sio populneo probably originates from the The tree is often planted as an ornamental, Asian tropics or from the coasts of the Pacific shade or roadside tree in coastal zones. In and Indian Oceans. It now occurs throughout Madagascar it has been planted to support the tropics in coastal areas; it is planted in vanilla vines. In India and on Pacific islands coastal towns and occasionaUy inland the tree is planted as a live fence and along the Uses The wood (trade names: milo, Seychelles coast as protection against erosion. In Asian rosewood, Polynesia rosewood, Pacific rosewood) mangrove areas it is planted to consolidate is used for a wide range of purposes where ridges and bunds in an aqua-SIIvicultural sys- quality is more important than size. In Ghana tern for prawn production. Thespesio populneo the wood is used for canoe seats, paddles, car is a sacred tree in many parts of the Pacific, parts, wheelbarrows and domestic utensils. In where it has often been planted near temples East Africa the wood sawn in small planks is and is used in traditional ceremonies. used to repair fishing boats. In the Indo-Pacific In traditional medicine, the bark, root, leaves, region the wood is considered excellent for flowers and fruits are used to treat a range of carving and is widely used for bowls and ailments, including skin problems, dysentery, plates, clubs, paddles, agricultural implements, cholera, haemorrhoids, liver and gall bladder musical instruments, gunstocks, carts, wheels, problems, urethritis, gonorrhoea, rheumatism boats, tool handles, furniture, cabinet work, and high blood pressure. In Nigeria the bark utensils, jewellery and turnery. It is also con- and a lotion from the boiled bark are applied to sidered suitable for light construction, floorlrig, wounds, and the seed oil to skin infections. In wall panelling, interior trim, precision equip- Mauritius the fruit sap is applied to warts. The merit, toys and novelties, and pattern making. heartwood is used to treat PIeuritis, cholera, The wood can be used asfirewood. colic and fevers, and is considered carminative The bark yields a tough fibre used for cordage, The cooked fruit crushed in coconut oil pro- fishing lines, basketry, coffee bags and for vides a salve that is applied to the hair to kill lice. Properties The heartwood is reddish brown to dark brown, often with purple veining; it Is sharply differentiated from the I-2 cm wide sapwood, which is white to pale yellow or pale . pink, darkening on exposure. The grain is shallowly interlocked to wavy, texture medium to fine. The wood shows slight ribbon figure on quarter-sawn faces. Freshly cut wood has a rose-like smell o The density of the wood is about 770 kg/in3 at Q. 12% moisture content. The wood seasons well, and does not warp or check. Shrinkage rates o0 are 3.8% radial and 6.9% tangentialfrom green

o to oven dry. Movement in service is very low. The wood is strong and hard. At 12% moisture content, the modulus of rupture is 1/8 N/mm2, Thespesiopopulneo- wild Gridplonted modulus of elasticity 11,690 N/min2, and coin- 548 TIMBERSl pression parallel to grain 56 N/min2. The wood is easy to saw and work with hand and ina- chine tools. It turns well in both green and dry conditions, can be finished to an attractive pol- I * ish and paints well. The wood contains an oil " y. I"' which slows down drying of varnishes. It has .. medium to poor gluing properties. ~\ The heartwood is very durable, even when in contact with water or the ground. It is resistant to termites, but not to marine borers. The \ . sapwood is not susceptible to Lyetus borers I The heartwood contains several sesquiterpe- I

* noid quinones, including mansonone D and H, .,. thespone and thespesone, which are known to . .... Induce contact dermatitis, to inhibit tumour . . 7" \ , formation and to have antifungal properties. , t, . I , The heartwood and other plant parts contain , \ . gossypol I I The ultimate bast fibre cells are 1.0-3.5 mm ^

long, with a diameter of 9-20 F1m. The fruits I I \ yield 0.4% of a flavonoid colouring matter V Thespesin, a sesquiterpene dimer, is the yellow . dye compound and is mainly obtained from the fruit. The seeds contain 18-34% oil, with 43- 2 49% linoleic acid, 21-34% palmitic acid, 14- 33% oleic acid and 2-3% stearic acid. Thespe- Thespesiopopu!neo- I, nowering bronch, . 2, fruit. sin, ceryl alcohol and ^-sitosterol have been Source. .PROSEA isolated from the unsaponifiable fraction of the seed oil main veins yellow, mostly with saccate nectar- The bark contains up to 7% tannin. Aqueous Ies in the axils of the basal veins beneath. and methanolic extracts of the bark have Flowers solitary in leaf axils, bisexual; pedicel shown in-vivo anti-oxidant activity in rats. The 1.5-10 cm long, erect or ascending; bypan- fruits and leaves contain compounds with anti- thium discoid, 6-8 mm in diameter; epicalyx bacterial activity, whereas methanolic extracts segments 3, oblong to lanceolate, 2-17 mm x 2 of the flower buds have shown antifungal activ- mm, early caducous, acute; calyx campanulate, ity. Ethanol extracts of the flower have shown 7-15 min long, truncate or slightly toothed at antihepatotoxic activity. Aqueous extracts of apex, densely appressed hairy inside, glabres- the fruit have shown wound-healing activity in cent outside; petals 5, obliquely obovate, 4-8.5 rats after topical or oral administration. The cm x 3.5-6 cm, apex rounded, pale yellow with seeds have purgative properties. The seed oil dark purple basis, scaly outside, glabrous in- has anti-amoebic activity. The root is toxic. side; stamens numerous, fused into a staminal Description Evergreen shrub or smalltree column, with free filaments 3-5 min long, an- up to LOG20) in tall; hole branchless for up to 3 thers c. 1.5 mm long; ovary superior, globose to in, often twisted or bent, becoming hollow with ovoid, 8-10 mm in diameter, scaly, 10-celled, age, up to 60(-120) cm in diameter, without style c. 4 cm long, stigmas club-shaped, pale buttresses; outer bark becoming rough and yellow. Fruit a depressed globose capsule 2-4.5 fissured in older trees, greyish, inner bark very cm in diameter, faintly 5-angular, apex obtuse fibrous, pink; crown broad and dense; twigs or srightly depressed, yellowish to brownish densely covered with minute brown to silvery green when mature, scaly, usually indehiscent, scales, glabrescent. Leaves arranged spiralIy, exuding a bright yellow gum when cut, many- simple and entire; stipules lanceolate to subu- seeded. Seeds obovoid, 8-15 mm x 6-9 min, late, 3-10 mm long; petiole 2-Ile16) cm long, slightly angular, covered by closely matted scaly; blade orbicular, deltoid, ovate or oblong, silky hairs. Seedling with epigeal germination; 6-23 cm x 5-16 cm, base cordate, apex acumi- hypocotyl elongated; cotyledons leafy; allleaves nate, rather fleshy and shiny, both surfaces arranged spiralIy. covered with small scales, palmately 7-veined, Other botanical information Thespesio

. THESPESiA 549 comprises about 15 species and is distributed parenchyma: 76: axial parenchyma diffuse; (77: throughout the tropics. Within Thespesio axial parenchyma diffuse-in-aggregates); (79: popu!neo, some authors recognize the speci- axial parenchyma vasicentric); 86: axial paren- mens occurring along the coasts of the Indian chyma in narrow bands or lines up to three Ocean as a distinct species: ritespesto popul- cells wide; 90: fusiform parenchyma cells; 91: neoides (Roxb. ) Kostel. , having somewhat two cells per parenchyma strand; (92: four (3- bronzed or coppery, shallowly cordate leaves, 4) cells per parenchyma strand). Rays: (97: ray pedicels 5-12 cm long and fruits with a dehisc- width I-3 cells); (98: larger rays commonly 4- ing outer layer. However, many intermediate to 10'seriate); 106: body ray cells procumbent specimens (called 'hybrids' by some) exist with one row of upright and/or square marginal where both types can be found. In Sri Lanka cells; 107: body ray cells procumbent with some of these 'hybrids' have been widely mostly 2-4 rows of upright and/or square mar- propagated vegetativeIy as ornamentals and ginal cells; 110: sheath cells present; (1/1: tile living fences cells present); 1/5: 4-12 rays per mm. Stoned Thespesio ocutilobo (Baker f. ) Exell & Men- structure: 1/9:low rays stoned, high rays non- donea ('wild tulip tree' or 'small-leaves tulip stoned; 120: axial parenchyma and/or vessel tree') is a shrub or smalltree up to 5(-6) in tall, elements stoned. Mineral inclusions: 136: occurring in Mozambique and Natal(South prismatic crystals present; 137: prismatic crys- Africa) in woodland and thickets on recent tals in upright and/or square ray cells; (141: sands near the coast. Its heartwood, which is prismatic crystals in non-chambered axial pa- dark and becomes hard and durable when sea- renchyma cells). soned, has been used for carving and for inak- re. Mugabi, A. A. Oteng-Amoako & P. Baas) ing spears, sticks and musical Instruments. A Growth and development Germination decoction of the bark is taken against chronic begins 8 days after sowing and may extend for dysentery. It is easy to cultivate as long as 10 weeks. Growth in height is rapid Thespesio donis 011v. is a shrub or smalltree in the first few years (0.5-1.5 in per year), but up to 6(-10) in tall, occurring in Ethiopia, So- slows down at 7-10 years of age. Stem diame- malla, Kenya and Tanzania in forest, bushland ter growth is I-3 cm/year. Flowering may al- and grassland up to 500 in altitude. The stems ready begin when the tree is only I-2 years are made into throwing-clubs, bows and ar- old. In southern Africa flowering is in Febru- rows, and are used as fire-sticks. The bark is ary-March and fruiting in MarchJune, in used for tying. The fruit is recorded to be edi- more even equatorial chinates flowering is ble. A dye is obtained from the flowers and year-round. The pale yellow flowers open at fruits. Root decoctions are taken against gonor- about 10 a. in. , turn reddish orange in the af- rhoea, stomach pain, pain in the spinal region, ternoon, fade to pink on the tree and do not fall haematuria and swelling of the abdomen off for several days. Pollination is probably by Anatomy Wood-anatointcal description OAWA birds. The seeds float and remain viable in hardwood codes): seawater for more than a year, making natural Growth rings: (I: growth ring boundaries dis- distribution by sea currents possible. Thespesio tinct); (2: growth ring boundaries indistinct or populneo naturalizes easily and has become a absent). Vessels: 5: wood diffuse-porous; 13: weed in some regions simple perforation plates; 22: intervessel pits Ecology 711espesiopopu!neo is a tree oftropi- alternate; (23: shape of alternate pits polygo- cal and warm subtropical climates and is usu- na1); 25: intervessel pits small (4-7 grn); (26: ally found up to 150 in altitude. The mean an- intervessel pits medium (7-10 pin)); 30: vessel- nualtemperature may range from 20-26'C, ray pits with distinct borders; similar to in- the mean annual rainfallfrom 500-5000 min tervessel pits in size and shape throughout the Occasional very light frost is tolerated. The ray cell; 41: mean tangential diameter of vessel tree grows best in full sunlight and does not Iumina 50-100 prn; 42: mean tangentialdiame- grow well in the shade of other trees. It toIer- ter of vessellumina 100-200 pin; 47: 5-20 ves- ates wind and salt spray. It is suitable for dry sels per square minimetre; 58: gums and other locations because it develops a long taproot in deposits in heartwood vessels. Tracheids and porous soils; It may tolerate a dry season of up fibres: 61: fibres with simple to minutely bor- to 8 months. 771espesio popu!neo thrives on dered pits; 63: fibre pits common in both radial sandy coastal soils as well as volcanic, lime- and tangential walls; 66: non-septate fibres stone and rocky soils with a pH of 6.0-7.4. It present; 69: fibres thin- to thick-walled. Axial tolerates heavier soils, soil sannity and occa- 550 TIMBERSl

SIonalinundation, but does not grow on per- genetic erosion inanently inundated soils. Natural stands oc- Breeding Although widely grown as an or- cur at the inland edges of mangrove swamps namental, named cultivars have not been re- and along tidal waters leased Propagation and planting Thespesio po- Prospects itespesiopopu!neo is a multipur- pulneo is usually propagated by seed, but pose tree suitable for dry and saline conditions, propagation by stem or root cuttings or by air- of which the wood is mainly used locally, e. g layering is also possible. The 1000-seed weight for carving. It is unlikely that the importance is 140-285 g. Seed storage behaviour is ortho- for sawn timber will Increase, but its local use dox, retaining viabinty when dried and stored. will remain. It may become increasingly impor- Germination can be difficult due to the hard tant as an ornamental and it is eminently suit- seed coat, and is improved by scarification with able for coastal erosion control. It should not be a knife, sandpaper or with concentrated SUI- promoted in cotton-growing areas, as it is a phuric acid for 20-60 minutes. Direct sowing is host of several cotton pests. generally practised. For plants grown in a Major references Abbiw, 1990; Balu Peru- nursery, pots must be large enough to accom- ina1, 1998; Burki11, 1997; Chowdhury & Ghosh, inodate the taproot. Seeds may be pre- 1958; Friday & Okan0, 2006; Keating & Bolza, germinated before being planted in pots. The 1982; Latiff & Fandah Hanum, 1997; Marais & plants are normally ready for planting out In Friedmann, 1987; Thulin, 1999a; Vonesen, 12-16 weeks, but trees up to 3.5 in tall have 1995a been planted out successfully from containers Other references Beentje, 1994; CAB In- Stump planting Involves cutting back the stem ternationa1, 2005; Coates Palgrave, 1983; to about I cm above the root collar before CSIR, 1972; Datta & Saha, 1970; Exell & transplanting, thus allowing the roots to re- Meeuse, 1961; Fosberg & Sachet, 1972; Grace, cover before new leaves develop. Wildlings are Ewart & Tome, 1996; Gunb-Fakim & Brendler, also collected and transplanted. For vegetative 2004; Gunb-Fakim, Gu6ho & Bissoondoyal, propagation small cuttings should be rooted in 1996; Hochreutiner, 1955; navarasan at al. , a nursery before planting out, but cuttings up 2003; InsideWood, undated; Kader & Chacko, to 2 in long have also been successfully planted 2000; Milbrodt, K6nig & Hausen, 1997; Na- directly in the field. gappa & Binu Cheriyan, 2001; Natarajan at al. , Management Weeding is important until 2005; Neuwinger, 2000; Shirwaikar at al. , the tree has become established. Thespesio 1995; Williams, 1949; World Agroforestry Gen- popu!neo tolerates heavy pruning, but re- tre, undated growth is slow. Nevertheless, trees should be Sources of illustration Latiff & Fandah pruned to develop clear holes for timber pro- Hanum, 1997 duction. It may take 25-40 years before the Authors P. Oudhia tree Is large enough to produce usable timber, although branches as little as 5 cm thick are used for carving. TIEGHEMELLAAFRICANA Pierre Diseases and pests Thespesio populneo is prone to root and stem rot caused by the fun- Protologue Not. hat' I: 18 (1890) gus Phellinus norius. Symptoms are slowly Family Sapotaceae enlarging diseased patches and a thick, dark Chromosome number2n=26 brown mycellal sheath around the base of in- Synonyms Dumorio afi. icono relerre) Dubard fected trees. It is also recorded to be susceptible (1915), Mimusops of neono (Pierre) Lecomte to fungal leaf spot (Lophodermium sp. ), heart (1921), Banjone110 o17icono (Pierre) Baehni rot (Fomespochyphloeus) and branch canker of (1965) tea (Phomopsis theoe) Vernacular names Douka (En). Douka (Fr) The tree is a host of a number of serious pests Origin and geographic distribution Tieghe- of cotton, such as cotton stainers (Dysdercus menu of neono occurs in Gamero0n, Equatorial spp. ), spiny bonworms (Bonus spp. ) and the Guinea, Gabon, Congo and DR Congo. How- bon weevil(Arithonomus grandis). Therefore, ever, some specimens have been reported from planting is discouraged or even outlawed in Cote d'Ivoire, where the closely related Tieghe- some cotton-growing areas. me!IQ hechem (Achev. ) Roberty occurs Genetic resources Thespesio populneu has Uses The wood, traded as douka, but often also a wide distribution and is not threatened by as makore or cherry mahogany (often without

. TIEGHEMELLA 551

shrinkage of the wood from 90% to 60% air moisture content was 1.1% radial and 1.8% tangential. Shrinkage from green to 12% ino1s- ture content was 3.0% radial and 4.5% tangential, and from green to oven dry 5.0-6.0% ra- . dial and 7.2-7.7% tangential. Usually, drying does not cause problems, although the wood dries slowly At 12% moisture content, the modulus of rup- ,^ ture is 96-138 N/min2, modulus of elasticity 10,100-13,850 N/min2, compression parallel to L, grain 51-59 N/min2, shear 12.6 N/min2, cleav-

.. age 27 N/mm and Janka side hardness 4940 N The wood is somewhat difficult to work due to

o the presence of silica; blunting effects are inod- erately severe when sawing, and stellite-tipped sawteeth are recommended. The wood finishes negheme!IQ of neono- wild well. Staining and polishing give good results Pre-boring for nailing and screwing is recoin- distinction from neghemello heekelii) , is used mended to avoid splitting. Gluing properties for furniture, exterior and Interior construction, are good. The timber can be peeled satisfacto- flooring, doors, vehicle frames, sports goods, rily railway sleepers, ship building, turnery and The heartwood is rated as one of the most du- sculptures, and makes good and decorative rable African timbers. It is resistant to ter- veneer, often used to face plywood. mites and fungi. Damage by pinhole borers and The seed kernels are rich in an edible fat, powder-post beetles has been recorded occa- which is locally popular as a cooking or season- SIonally. Although it is resistant to marine ing oil. In Gabon the fat is also used externalIy borers in temperate waters, the wood is not to treat rheumatism very durable in tropical (especiaUy brackish) Production and international trade Douka waters. The heartwood is very difficult to jin- is traded on the international timber market, pregnate, the sapwood moderately easy but production is small due to limited supply Dust from sawn wood may cause irritation to from naturalstands. It is often traded together skin and mucous membranes. It has been sug- with makore timber (from neghemello heeke- gested that this is caused by the presence of lit). The export from Gabon increased from saponins or the contact allergen 2.6-dimethoxy- 15,600 in3 in 1997 to 36,000 ing in 2001, but 1.4-henzoquinone. A highly haemolytic saponin decreased again to 25,000 ina in 2003. The ex- has been isolated from the wood; on hydrolysis port from Garnero0n is much lower: 390 in3 in it gave d-glucose, I-rhamnose and d-xylose. The 2003. In 1994 the price of sawn douka wood resistance of the wood to termites has been from Gabonwas Us$ 931ma. demonstrated in toxicity tests of extracts. Properties The data on wood properties of The fatty oil from the seed kernelis yellowish douka and makore cannot be separated in the and semi-fluid, has a pronounced flavour and literature, and the following description applies taste, and consists of about 55% oleic acid, to both species. The wood resembles African 21.5% stearic acid, 16.5% palmitic acid and 5% mahogany (Khoyo and Errtondrophrogmu linoleic acid. The kernel comprises about 50% spp. ), but the texture is finer. The heartwood is offat pinkish-, purplish- or reddish-brown with a Adulterations and substitutes Douka is silky lustre, often with a decorative figure in used for similar purposes as African mahogany the form of flames or stripes; it is very dis- (Khoyo and Errtondrophrogmo spp. ), but it is tinctly demarcated from the up to 8 cm wide more durable. It is sometimes even traded as and pinkish white to greyish brown sapwood African mahogany. It is very similar to the The grain is straight or interlocked, texture wood of makore (negheme!IQ heekelii), which is fine and even often traded together with douka under the The wood is medium-weight with a density of same commercial name of makore 600-800 kg/in3 at 12% moisture content. The Description Very large tree up to 55 in tall; shrinkage rates are low to moderate. In a test, hole up to 250 cm in diameter, sometimes much 552 TIMBERSl

penor, conical, softly hairy, 8-celled, each cell .:* ,' with I ovule, style short. Fruit a large, ovoid, .. , smooth berry c. 8 cm long, brownish yellow when ripe, containing I-3 seeds in a yellowish 3 pulp. Seeds broadly ellipsoid or ovoid, slightly lateralIy compressed, 5-7 cm long, testa thick, woody, smooth, shining and brown in dorsal part, rough and bullate in ventral part (scar); endosperm absent. Seedling with epigeal germination; hypocotyl stout, up to 17 cm long, epicotyl up to 17 cm long; cotyledons thick, sessile, c. 4 cm x I cm, green Other botanical information The genus neghemello consists of 2 species and is closely related to Mimusops, which differs in having * less-developed corolla tubes, smaller fruits, and seeds with smaller scars, copious endosperm ,,.., JJ ally I 2 I and thin cotyledons. Tiegheme!!o heekelii from West Africa resembles neghemello of neono ,- I " I and may be conspecific. The former differs in the smaller median segment of the corolla lobes, smaller staininodes and larger seed scar \ A taxonomic study is needed to clarify the spe- '!'''''''it, ,;,*.*"*~*$*;;\," 11;; 5 ..,. , 4 cies limits and the status of the genus, which is complicated by the fact that the name neghe- neghemello of neon0 - I, 60se of 601e, . 2, flower- me110 was first published for a genus offungi ing twig, 3, flower, 4, fruit, ' 5, seed. Anatomy Wood-anatomical description OAWA Redro"it und oddpted by AChmod Sottri Nur- hardwood codes) homerit Growth rings: 2: growth ring boundaries iridis- tinct or absent. Vessels: 5: wood diffuse-porous; more, straight and cylindrical, often swollen in 7: vessels in diagonal and/or radial pattern; 13 lower part, reaching up to 30 in to the first simple perforation plates; 22: intervessel pits branches, sometimes with buttresses; bark alternate; (23: shape of alternate pits polygo- surface brown to red-brown, deeply furrowed na1); 26: intervesselpits medium (7-10 pin); 30 with rectangular scales, inner bark fibrous but vessel-ray pits with distinct borders; similar to brittle, exuding a sticky latex; crown heavy, intervessel pits in size and shape throughout rounded, heavy branches often abruptly the ray cell; 31: vessel-ray pits with much re- spreading. Leaves arranged spiralIy, more or duced borders to apparently simple: pits less in tufts at the ends of branches, simple; rounded or angular; 32: vessel-ray pits with stipules absent; petiole 1.5-3.5 cm long, SIen- much reduced borders to apparently simple der; blade elliptical to obovate, 8-16 cm x 4-7 pits horizontal(scalariform, gash-like) to verti- cm, curieate at base, rounded or obtusely acu- cal (palisade); (33: vessel-ray pits of two dis- inmate at apex, with entire to slightly undulate tinct sizes or types in the same ray cell); 42 margin, thinly leathery, glabrous, lateral veins mean tangential diameter of vessellumina numerous. Flowers in fascicles of 2-3 in the 100-200 pin; 47: 5-20 vessels per square inn- leaf axils, bisexual, regular; pedice1 1.5-2.5 cm limetre; 56: tyloses common; (58: gums and long; calyx with 2 whorls of 4 lobes c. 6 min other deposits in heartwood vessels). Tracheids long, outer ones glabrous but softly hairy at and fibres: 61: fibres with simple to minutely margins, inner ones softly hairy outside; co- bordered pits; 66: non-septate fibres present; rolla with c. 1.5 min long, fleshy tube and 8 69: fibres thin- to thick-walled; (70: fibres very lobes, each lobe with Ifiliform median segment thick-walled). Axial parenchyma: 86: axial pa- and 2 large, fleshy, jinbricate lateral segments Tenchyma in narrow bands or lines up to three c. 4 mm long, creamy white; stamens 8, in- cells wide; 87: axial parenchyma reticulate; 92: serted on the corolla tube in front of the corolla four (3-4) cells per parenchyma strand; 93: lobes, free, filaments short, 8 longer staini- eight (5-8) cells per parenchyma strand; 94 nodes alternating with the stamens; ovary su- over eight cells per parenchyma strand. Rays

. TIEGHEMELLA 553

97: ray width I-3 cells; 107: body ray cells pro- genetic erosion than neghemel!a heeke!it. It cumbent with mostly 2-4 rows of upright suffers less from habitat destruction than its and/or square marginal cells; 108: body ray West African relative, but is subject to selective cells procumbent with over 4 rows of upright logging in many regions and/or square marginal cells; 1/5: 4-12 rays Prospects The prospects for planting proper mm; (1/6: Z 12 rays per mm). Mineral in- grammes using douka are good in the light of clusions: 159: silica bodies present; 160: silica the experiences in Gabon and with makore in bodies in ray cells West Africa. Like the latter species it may be (L. N. Banak, H. Beeckman & P. E. Gasson) suitable for use in agroforestry programmes, Growth and development Young douka being not only important for its timber, but trees were on average 9 in tall 6 years after also for fat production from the seeds. planting in Gabon. The survival rate was about Major references Aubr6ville, 1964; CIRAD 90% and was almost equal when planted ex- Forestry Department, 2003; de Saint-Aubin, posed to fullsunlight in comparison with plant- 1963; Koumba Zaou at a1. , 1998; Louppe at al. , ing in the shade of forest cleared from under- 1999; Raponda-Walker & Sillans, 1961; Richter growth. The young trees grow faster when & Danwitz, 2000; Wilks & Issemb6, 2000 planted in light shade; 11-year-old trees were Other references Aubr6ville, 1961; Bile N- on average 18.5 in tall and 13 cm In diameter logh0, 1999; Bokdam, 1977; Gassagne, 1966; when planted in light shade, and 15.5 in tall Gassita at al. (Editors), 1982; InsideWood, un- and 9 cm in diameter in full sunlight. In a 66- dated; Takahashi, 1978; Vivien & Faure, 1988c year-old plantation in Gabon the mean annual Sources of illustration Aubr6ville, 1964; increment is 0.4 cm in diameter and 1.8 ingftia Wilks & TSSemb6, 2000 in wood volume. The fruits are eaten by ele- Authors R. H. M. J. Lemmens phants, which are probably the main seed dispersers Ecology Douka is an emergenttree of primary TIEGHEMELLAHECKELli(Achev. ) Roberty rain forest. It is most common in coastal lowland and diminishes towards the eastern parts Protologue Petite F10re de 1'0uest-Africain: of Cameroon, Equatorial Guinea and Gabon. It 79 (1954). occurs scattered in the forest, but may locally Family Sapotaceae be more abundant. Young trees are tolerant of Chromosome number2n=24 shade, but can survive in fullsun Synonyms Dumorio heekeliiA. Chev. (1907), Propagation and planting The seeds are Mimusops heekelii(Achev. ) Hutch. & Dalziel heavy: 15-20 g. They should be planted within (1931), Boillone!10 hechant (Achev. ) Baehni a few weeks because viahinty decreases rapldly. (1965). Management The density of douka in the Vernacular names Makore (En). Makor6 forest is generally low. The average of 14 in- (Fr). Makor6 (Po). ventories in western Gabon was 0.5 ms oftim- Origin and geographic distribution neghe- her per ha. The total volume of douka timber in Gabon was estimated at 6 million ina. The minimum diameter allowed for exploitation is 70 cm in Gabon and 60 cm in Garnero0n.

Douka has been planted on a smallsca}e (37 ha . in 1988 and 1989) near Ekouk (Gabon), and still smaller plantations exist elsewhere in Gabon, where the results indicate that douka is one of the best indigenous species for planting after okoum6 (Aucoumeo hloineono Pierre) ,^

Handling after harvest Care should be L. taken when sawing douka wood. Nasal and

respiratory Irritation with haemoptysis oc- .. curred in men sawing wood from Equatorial Guinea. o Genetic resources Although neghemello of- ricono is also included in the IUCN red list of threatened species, it is probably less liable to neghemello heekelii - wild

. 554 TIMBERSl

meno heckelii occurs in the West African forest tangential. Shrinkage from green to 12% inois- zone, from Sierra Leone eastwards to southern ture content was 3.0% radial and 4.5% tangen- Nigeria, but not in Togo or Benin tial, and from green to oven dry 5.3-6.5% ra- Uses The wood, traded as makore or cherry dial and 7.3-8.7% tangential. Usually, drying mahogany, is used for furniture, exterior and does not cause problems, although the wood interior construction, flooring, doors, vehicle dries slowly. frames, sports goods, railway sleepers, turnery At 12% moisture content, the modulus of rup- and sculptures, and makes good and decorative ture is 96-165 N/min2, modulus of elasticity veneer, often used to face plywood, especially 9500-13,900 N/mm2, compression parallel to for marine uses grain 47-67 N/min2, shear 8-14 N/mm2 cleav- The seed kernels (cotyledons, known as 'baco') age 17-27 N/mm and Janka side hardness are rich in an edible fat known as 'dumori but- 4940 N ter' or 'makore butter', which is locally popular The wood is somewhat difficult to work due to as a cooking or seasoning oil and often pre- the presence of silica; blunting effects are inod- forred to palm oil. The fat is also applied as a erately severe when sawing, and stellite-tipped pomade to the body and hair, and used in soap sawteeth are recommended. The wood finishes production. The fleshy and very sticky pulp of well. Staining and polishing give good results the fruits is sometimes used as birdlime. The Pre-boring for nailing and screwing is recoin- bark is reportedIy effective for treating blen- mended to avoid splitting. Gluing properties norrhoea and toothache, and in Liberia young are good. The timber can be peeled satisfacto- buds are used to treat snake bites rily Production and international trade Makore The heartwood is rated as one of the most du. Is traded on the international timber market, rable African timbers. It is resistant to ter- but production is small due to limited supply mites and fungi. Damage by pinhole borers and from naturalstands. It is often traded together powder-post beetles has been recorded DCca- with 'douka' wood (from Tiegheme110 o17icuno SIonally. Although it is resistant to marine Pierre). The export of makore timber from C6te borers in temperate waters, the wood is not d'Ivoire declined from 70,000 in3 in 1960 to very durable in tropical (especially brackish) 28,000 in3 at the beginning of the 1980s and waters. The heartwood is very difficult to jin- 6000 ms at the end of the 1980s. In 1994 pregnate, the sapwood moderately easy Ghana exported 2100 ina of air. dried makore Dust from sawn wood may cause irritation to sawnwood for an average price of Us$ 5101m3, skin and mucous membranes. It has been sug- and 3200 in3 of sliced veneer for an average gested that this is caused by the presence of price of Us$ 7801m3, rotary-peeled veneer for saponins or the contact allergen 2.6-dimethoxy- Us$ 4501m3 and jointed veneer for Us$ 1.4-henzoquinone. A highly haemolytic saponin 1735/ina. C6te d'TVoire exported 200 ing of ve- has been isolated from the wood; on hydrolysis neer for Us$ 1800/ina in 1994. In 2001 Ghana it gave d-glucose, I-rhamnose and d-xylose exported 3500 ms of makore veneer The fruit is sticky and juicy, with an unpleas- Properties The data on wood properties of ant smell and bitter taste. The kernel coin- makore and douka cannot be separated in the prises about 60% oil by weight. The o111s yel- literature, and the following description applies lowish and semi-fluid, has no distinct flavour to both species. The wood resembles African or taste (occasionally slightly spicy), and con- mahogany (Khoyo and Errtondrophrogmo SISts of about 51% oleic acid, 43% stearic acid, spp. ), but the texture is finer. The heartwood is 3.5% palmitic acid and 2.5% linoleic acid pinkish-, purplish- or reddish-brown with a Adulterations and substitutes Makore is silky lustre, often with a decorative figure in used for similar purposes as African mahogany the form of names or stripes; it is very dis- (Khoyu and Errtondrophrogmo spp. ), but it is tinctly demarcated from the up to 8 cm wide more durable. It is sometimes even traded as and pinkish white to greyish brown sapwood. African mahogany. It is very similar to the The grain is straight or interlocked, texture wood of Tieghemello of neono from Central fine and even Africa, which is often also traded as makore or The wood is medium-weight with a density of douka 600-800 kg/ing at 12% moisture content. The Description Very large tree up to 55 in tall; shrinkage rates are low to moderate. In a test, hole up to 250 cm in diameter, sometimes much shrinkage of the wood from 90% to 60% air more, straight and cylindrical, often swollen in moisture content was 1.1% radial and 1.8% lower part, reaching up to 30 in to the first

. TIEGHEMELiA 555

conical, softly hairy, 8-celled, each cell with I ovule, style short. Fruit a large, ovoid-globose,

.\ smooth berry 8-12 cm long, yellow when ripe, containing I-3 seeds in a yellowish pulp. Seeds \.;#,, 1" broadly ellipsoid, slightly lateralIy compressed, 2 (596-7.5 cm long, testa thick, woody, smooth, \I shining and yellowish-brown in dorsal part, "'., <:1^ "' 11 rough and bullate in ventral part (scar); en- If'\ I 1<*' dosperm absent. Seedling with epigeal germi- \ .. L-- nation, hypocotyl stout, 6-14 cm long, epicotyl

I::: ^-.~ ,,. ., up to 3-11 cm long, cotyledons thick, sessile, c I' .I \ -.^. ^. *\ 5 cm x 2 cm, dark green Other botanical information The genus

., I' Tieghemello consists of 2 species and is closely I related to Mimusops, which differs in having less-developed corolla tubes, smaller fruits, and seeds with smaller scars, COPIOUS endosperm and thin cotyledons. Tieghemello of neono re- \.; \ sembles neghemello heekelii and may be con- If'\' .. specific. The former differs in the longer, fill-

I' form median segment of the corolla lobes, Iar- 11/11 ^i ger staininodes and slightly smaller seed scar 9\ A taxonomic study is needed to clarify the spe- \^"' ',I' 11;;;\. \ I I cies limits and the status of the genus, which is 4 5 3 "-111.7 I complicated by the fact that the name Tieghe- menu was first published for a genus offungi. neghemello heekelii- I, flowering twig, . 2, flower, Anatomy Wood-anatomical description OAWA 3, fruit, 4, seed, . 5, seed kernel hardwood codes) Redrawn ond adopted by W. We88e!-Byund Growth rings: a growth ring boundaries iridis- tinct or absent. Vessels: 5: wood diffuse-porous; branches, angular or ridged at base, or with 7: vessels in diagonal and/or radial pattern; large, narrow buttresses up to 3 in high and (10: vessels in radial multiples of 4 or more heavy, spreading surface roots; bark c. 2 cm common); 13: simple perforation plates; 22: thick, surface grey to nearly black, deeply fur- intervessel pits alternate; 23: shape of alter- rowed with rectangular scales, inner bark red- nate pits polygonal; 25: intervessel pits small dish, fibrous but brittle, exuding a sticky latex; (4-7 F1m); 26: intervessel pits medium (7-10 crown heavy, rounded, heavy branches often Urn); (30: vessel-ray pits with distinct borders; abruptly spreading. Leaves arranged spiralIy, similar to intervessel pits ln size and shape more or less in tufts at the ends of branches, throughout the ray cell); 31: vessel-ray pits simple; stipules absent; petiole 1.5-4 cm long, with much reduced borders to apparently SIm- slender; blade elliptical to obovate, 6-15 cm x PIe: pits rounded or angular; 32: vessel-ray pits 2-6.5 cm, curieate at base, emarginate, with much reduced borders to apparently SIm- rounded, acute or acuminate at apex, with en- PIe: pits horizontal (scalariform, gash-like) to tire to slightly undulate margin, papery or vertical (palisade); (33: vessel-ray pits of two thinly leathery, glabrous, lateral veins numer- distinct sizes or types in the same ray cell); 42 ous. Flowers in fascicles of I-4 (as a rule 2) in mean tangential diameter of vessellumina the leaf axils, bisexual, regular; pedice1 1.5-2.5 100-200 prn; 47: 5-20 vessels per square inn- cm long; calyx with 2 whorls of 4 lobes 3-5 min limetre; 56: tyloses common; (58: gums and long, softly hairy on the overlapplrig parts; other deposits in heartwood vessels). Tracheids corolla with 2-2.5 min long, fleshy tube and 8 and fibres: 61: fibres with simple to minutely lobes, each lobe with I minute median segment bordered pits; 66: non-septate fibres present; and 2 large, fleshy, jinbricate lateral segments 69: fibres thin- to thick-walled. Axial paren- 3-3.5 min long, creamy white; stamens 8, in- chyma: 86: axial parenchyma in narrow bands serted on the corolla tube in front of the corolla or lines up to three cells wide; 87: axial paren- lobes, free, filaments short, 8 short staininodes chyma Teticulate; 93: eight (5-8) cells per pa- alternating with the stamens; ovary superior, renchyma strand; 94: over eight cells per pa-

. 556 TIMBERSl

Tenchyma strand. Rays: 97: ray width I-3 cells; are very tolerant of shade, but can survive in 100: rays with multiseriate portion(s) as wide full sun. Heavy soils are preferred. as uniseriate portions; 107: body ray cells pro- Propagation and planting Makore is easy cumbent with mostly 24 rows of upright to propagate. The seeds are heavy, 30-50 and/or square marginal cells; 108: body ray seeds/kg. Viahinty decreases rapidly. They cells procumbent with over 4 rows of upright should be planted within 2 weeks. In the nurs- and/or square marginal cells; 1/5: 4-12 rays ery, seeds are usually planted at a spacing of per mm. Mineral inclusions: 159: silica bodies 40 cm x 40 cm under light shade. Seedbeds present; 160: silica bodies in ray cells should be watered well. Germination is over (L. N. Banak, P. 06tienne & P. E. Gasson) 90% in optimal conditions, and without pre- Growth and development Initial growth treatment. It begins after about 4 weeks, most of seedlings is fast, up to 70 cm in 4 months, seeds having germinated after 10 weeks, but it but then often ceases while a strong taproot (c may continue for 16 weeks. Rodents may cause 20 cm long)is formed. Growth of young makore considerable damage to seeds and seedlings trees is reported as slow, but strongly depends (cotyledons). Saplings of about 2.5 years old on light. Under conditions above 40% of full and I-1.5 in tall are planted out into the field light, growth may be up to I in/year; under usually at the beginning of the rainy season, 10% growth is almost nil. Other reports indi- but in western C6te d'TVoire 0.5 year-old seed- cate, however, that maximum growth is lings are used reached at an Irradiance of 10% of unshaded Propagation by air layering and cuttings is values. In Ghana, young trees have been re- successful. In a test with air layering, 39% of ported to be I-2 in tall after I-1.5 years in a branches rooted after 8 weeks. The average nursery, and when planted under shade trees rate of rooting of branch cuttings after 6 in the field 1.5-3.5 in tall after 13 years. How- months was 73%. In neither case did the appli- ever, in western C6te d'TVoire, trees have been cation of indole-butyric acid Improve rooting reported to be 3 in tall after 4 years and 9-11 success in tall after 20 years with a stern diameter of Management The density of neghemello 13-16 cm, occasionally even 28 in tall after 21 heekelti in the forest is generally low. Reports years with a diameter of 37 cm. For 35-80- for C6te d'Ivoire vary from I exploitable tree year-old trees the mean annual increment is per 23 ha to 1.4 trees greater than 10 cm di- 0.4-0.8 cm ameter per ha. The average of 9 inventories in Trees develop according to Aubr6ville's model: C6te d'TVoire was 4.1 in3 of timber per 100 ha the monopodialtrunk shows rhythmic growth, for trees exceeding 70 cm diameter. The aver- with whorled branches, which also grow age of 16 inventories in Liberia was 33 in3 of rhythmicalIy but modularly, each branch PIa- timber per 100 ha for trees exceeding 40 cm giotropic by apposition, the modules growing diameter. Experiments in C6te d'Ivoire showed indefinitely that thinning of the forest leads to better re- Trees start flowering and fruiting after about CTuitment and growth. 17 years, but sometimes after 10 years. Flow- Makore has been planted on a very smallscale ers open in the early morning, the corolla being (almost 6 ha), mainly for seed oil production, in shed in the afternoon of the same day. In Libe- western C6te d'Ivoire (near Tai National Park) ria, flowering is from February to May; ripe with an average density of 357 stemsftia. It is fruits can be found between October and De- often planted at a density of about 120 cember. In Cote d'Ivoire, flowering occurs from trees/ha, in association with coffee, cocoa, rub- January to June, and ripe fruits can be found ber or rice, and at field margins and in secon- from August to March. A large tree produces dary forest following clearance of undergrowth approximately 3000-4000 fruits at a time. The Diseases and pests No information on dis- fruits are eaten by elephants, which are proba- eases and pests is available, although neghe- bly the main seed dispersers; bush-pigs are menu heekeliiis a host of the semi-parasitic also reported to feed on the fruits tree Okouboho uubreuilleiPellegr. & Normand. Ecology neghemello heekeliiis often an Harvesting Logs tend to split badly during emergent tree of the high forest. It is found in felling. Logs of larger trees may be hollow moist evergreen and semi-deciduous forests. Yield Fat yield is 20-30 kg/tree per harvest. Regeneration in these forests is reportedIy rea- One kg of seed kernels yields about 200 g offat sonable, at least in Ghana; there is a strong when the traditional method of extraction is preference for undisturbed forest. Young trees used

. TooNA 557

Handling after harvest Care should be Vernacular names Toon, Indian mahogany, taken when sawing makore wood because na- Australian red cedar (En). C6dre rouge, c6dre sal and respiratory irritation may occur. To rouge d'AUStralie (Fr). Cedro australiano (Po). extract the oil, the seeds are split, and the ker- Origin and geographic distribution Toono nels dried in the sun, roasted, pounded to a enjoto originates from tropical Asia and tropi- paste and boiled in water. The fat or oilis cal Australia, but is now much cultivated skimmed off from the water throughout the tropics for Its timber and as an Genetic resources neghemello hecheliiis ornamental or wayside tree. It is extensively reported to be rather rare in Liberia. The den- planted in tropical Africa, particularly in East sity in C6te d'TVoire is low, and in many areas and southern Africa, but also locally in West it is considered rare. In Ghana, it is fairly Africa, Madagascar and Mauritius. It was re- common, but under pressure as a result of for- corded from Zambia and Zimbabwe as early as est exploitation, and deserves protection. The the beginning of the 20th century. It has locally seeds are dispersed by elephants, which are become naturalized in southern Africa becoming rare in West Africa and therefore Uses In South-East Asia the wood is consid- ITmiting natural regeneration ered of high value and used in house and ship neghemello heekeliiis included in the TUGN building, for joinery, high-grade furniture, tea Red list of threatened species because it suffers chests and boxes, musical instruments, toys both from habitat destruction and selective and novelties, carvings, veneer, plywood and logging practices pencils. Elsewhere the wood of planted Toono Prospects The prospects for planting pro- of!iota trees is occasionally used, also in tropi- grammes in West Africa using Tieghemello calAfrica heekelii are good. Seed and vegetative propaga- The flowers yield a reddish or yellowish dye, tion is easy. The species can be planted in open which has been used in tropical Asia to colour sites, grows fairly rapidly, and has a fairly silk. The bark may be used for tanning leather, open crown structure which permits good light and has been traditionally used to make twine penetration, making it suitable for use in agro- and string bags. Various parts of the plant, but forestry programmes. The wood, and fat from especially the bark, are used in traditional the seeds, are of excellent quality. However, medicine in the original distribution area of planting for fat production may be uneconomic Toono enjoto, e. g. as an astringent and tomc, as it takes a rather long time for trees to fruit to treat dysentery and to heal wounds. In Zim- after planting, and fat yields are limited babwe a leafinfusion is taken to treat venereal Planting is needed to take pressure off wild diseases. Toono ci!joto is commonly planted as populations an ornamental tree, and particularly as a road- Major references Ayensu & Bentum, 1974; side tree. This is the most common use in at- Bonn6hin, 2000; Burki11, 2000; CIRAD For- rica. It is also planted as a firebreak and for estry Department, 2003; CTFT, 1976a; Heme, reforestation. In Burundiit Is planted as a 1963; Richter & Danwitz, 2000; SIepel, Poorter shade tree in banana plantations and for ero- & Hawthorne, 2004; Voorhoeve, 1979 SIon control, and the sale of its timber is an Other references Agyeman, Swalne & important source of income for farmers. The Thornpson, 1999; Bokdam, 1977; Durrieu de foliage can serve as fodder, and has been used Madron at a1. , 1998a; Gosse at a1. , 2002; Haw- in tropical Asia as a vegetable. All aromatic oil thorne, 1995; InsideWood, undated; Takahashi, can be extracted from the wood and fruits 1978; Van Rompaey, 1993; Wiselius, 1994. Flowering trees are a good source of nectar for Sources of illustration Voorhoeve, 1979 honey bees Authors L. Bonn6hin & R. H. M. J. Lemmens Production and international trade In many regions within the natural area of distribution of Toono ciliato, the timber is highly TooNAciLiATAM. Roem. prized and has been much overexploited, particularly in Australia where it once was the Protologue Fam. nat. syn. monogr. I: 139 most important native timber. Nowadays it is (1846) exploited in many areas in South-East Asia, Family Menaceae e. g. in Myanmar. It Is exported in small Chromosome number 211= 52, 56, 78 amounts to China and Japan, but trade statis- Synonyms Cedrelo toono Roxb. ex Rottler & tics are not available Wind. (1803) Properties The heartwood is pale red to red-

. 558 TIMBERSl dish brown, darkening to dark red-brown on tresses at base; bark surface greyish white to exposure, usually distinctly demarcated from brown, usually fissured and flaking, inner bark the greyish white to pink sapwood. The grain is brown to reddish, fibrous; crown rounded, usually straight, sometimes interlocked, tex- spreading. Leaves alternate, paripinnately ture rather coarse and uneven. The wood has a compound with (5-)9-15 pairs of leaflets; stip- cedar-like odour. ules absent; petiole 6-11 cm long, rachis The wood is lightweight to medium-weight, slightly hairy or glabrous; petiolules 2-LOG14) with a density of 330-600 kg/ina at 12% inois- min long; leaflets lanceolate to ovate- ture content. The rates of shrinkage are usu- lanceolate, (7-)9-13(-16) cm x (2-)3-5(-6) cm, ally moderate. The wood may be liable to warp- asymmetric at base, acute or acuminate at ing and cupping during drying, particularly in apex, entire, glabrous, pinnately veined. Trillo- thin planks. Close spacing of stickers and rescence an axillary or terminal, much- weighting of stacks is recommended. Boards 25 branched, pendent panicle up to 55 cm long, min thick take I-3.5 months to air dry. Once hairy. Flowers unisexual, male and female dry, the wood is moderately stable in service flowers very similar in appearance, regular, 5- For wood of South African origin at 12% ino1s- merous, fragrant; pedice1 0.5-I mm long; calyx ture content, the modulus of rupture is 76 c. I mm long; petals free, 3.5-6 mm long, N/mm2, modulus of elasticity 8900 N/min2 creamy white; stamens free, I-2.5 mm long; compression parallel to grain 42 N/mm2, shear disk 1.5-2.5 min in diameter, reddish orange; 7 N/min2, Janka side hardness 3650 N and ovary superior, I-2 min in diameter, 5-celled, Janka end hardness 5330 N. style 0.5-3 min long, stigma head-shaped; male The wood is easy to saw, cross-cut and plane, flowers with rudimentary ovary, female flowers and the planed surface is smooth; it takes a with non-dehiscing arithers. Fruit an ellipsoid good polish. Some material tends to produce a to obovoid capsule 1.5-2.5 cm long, pendulous, woolly finish and the use of sharp tools is smooth or with lenticels, reddish brown, de- therefore recommended. Mortising, turning hiscmg with 5 slightly woody valves, many- and sanding give moderate results, boring seeded. Seeds I-2 cm long, winged at both sometimes poor results. Nailing Is easy, butthe ends with unequal wings. Seedling with nail-holding capacity is moderate. The gluing epigeal germination; cotyledons leaf-like; first properties are rated as good. The wood peels leaves opposite, 3-foliolate with lobed or welland the veneer is of good quality and has a toothed leaflets nice figure. The veneer can be glued to produce Toono cilioto grows rapidly. In Hawaii an av- good-quality plywood. erage tree height of 10 in and an average hole The wood is non-durable to moderately dura- diameter of 96 cm had been reached after 8.7 ble. It is usually susceptible to termite and dry- years. Mean annual diameter increment is wood borer attacks. The heartwood is usually 08-1.8(-2.5) cm. Trees may reach 35 in tall resistant to impregnation with preservatives, with a hole diameter of 70 cm when 40 years butthe sapwood is permeable. Wood dust may old. In plantations in tropical Africa, growth irritate the respiratory organs and skin. declines seriously after 40 years. Trees planted An ethanol extract of the heartwood showed in open localities may already flower and pro- anti-ulcer, gastro-protective and analgesic ac- duce seed after 6 years, The flowers are polli- tivities in tests with rats. Stem bark extracts nated by insects such as bees and moths. exhibited in-vitro antibacterial activity. Sev- Fruits ripen about 3 months after flowering erallimonoids have been isolated from Toono The seeds are dispersed by wind enjoto. The tetranortriterpenoid cedrelone Toono comprises 4 or 5 species and occurs in showed antifungal activity tropical Asia and eastern Australia. It is The foliage contains 13-14% crude protein and closely related to Cedre!o from tropical Airier- 14-22% crude fibre, and is reported to have ICa. Cedre!o odoroto L. is planted as a roadside good nutritive value, but poor palatability tree and shade tree in tropical Africa, also in Tests in Malawi showed that Toonu cilioto timber plantations, and has often been con- leaves were fairly palatable to sheep. Bark fused with Toono ciliato. It differs in its flow- extracts have insect-repellent activity ers having a longer column (androgynophore) Botany Deciduous or nearly evergreen, ino- on which the stamens and pistil are borne and noecious, medium-sized tree up to 25(-35) in in its entire seedling leaflets tall; hole branchless for up to 22 in, up to Toonu sinensis (A. Juss. ) MRoem. is planted in 50(-100) cm in diameter, with or without butsome African countries, e. g. Uganda and Tan-

. TooNA 559 zania, mainly as a roadside tree. It differs from many regions Toono enterto plantations are Toono enjoto in its toothed leaflets and seeds severely damaged by attacks of tip moth (Hyp- winged at only one end. The wood properties sipylo robusto), which may attack young are coinparable to those of Toonu dimtu shoots, flowers, fruits and seeds. Tip moth and Ecology In its natural area of distribution some other pests may cause seed losses of up to Toono enjoto occurs in primary as well as sec- 97%. Toono cilioto has been used successfully ondary forest, often along rivers and in valleys, in South-East Asia for enrichment planting, up to 1500 in altitude, in areas with 800-1800 and was then much less severely attacked by min annual rainfall. It is capable of regenera- tip moth than when seedlings were planted in tion in full sunlight. It prefers well-drained the open. The shoot borer Hypsjpylo grondel!o sites on deep, fertile soils, and does not grow attacks Toono ciliato worldwide. In the 1970s well in sandy localities. It tolerates some in several areas of Malawi many Tool, a echoto drought if the tree is well established. It is trees showed die-back of the branches, locally frost hardy. In Malawiit grows wellin areas at resulting in 80% mortality. It has been sug- 450-1500 in altitude with 900-1500 mm an- gested that Fusorium sp. was the causal organ- nualrainfall Ism. Management Trial timber plantations have Genetic resources and breeding Toono been established in Tanzania, Malawi, Zambia, ciliato is much sought after for its timber in Its Zimbabwe and South Africa. It has been re- natural distribution area, and has become ported from East Africa that Too, IQ emuto can scarce in many regions. It has been suggested be aggressive. It spreads easily by seed and cut that there is much genetic variation in Toono roots produce suckers. The root system spreads enterto over its large natural distribution area widely and develops close to the soil surface, Prospects Toono enjoto is useful for many and the use of the tree in gardens or crop plan- purposes. It has been recommended in AUStra- tations is dissuaded. Seeds are often produced 11a for planting because of its high-value tim- in great number, and can be collected from the her, in Mexico as a shade tree in coffee planta- soil. The 1000-seed weight is 2.5-5 g. Seeds can tions and in India as a multipurpose tree in be stored for only a few months at room tern- agroforestry systems. In tropical Africa the perature, but for seed stored at -4'C a viabillty results of plantation establishment were in of 97% has been recorded after 5 years and of general favourable, although plantations in 38% after 12.5 years. The seeds are best placed Sudan showed poor hole shapes and planta- in the ground with a wing pointing upward tions in some localities in Malawisuffered from The germination rate is 30-80%, and seeds serious die-back. The attractive multipurpose germinate 7-28 days after sowing. In Malawi wood and the fast growth make Too, IQ enjoto best germination results were obtained when attractive for planting. On the other hand, the fully mature seeds were sown in the nursery on superficial root system is a drawback for plant- soil collected in Brachystegio woodland. In In- ing in agroforestry systems, and its susceptibil- dia I-year-old seedlings are preferred for field ity to diseases and pests should be considered planting. In Australia placing plastic tubes when using this species in timber plantations. around seedlings has been recommended. Root The planting of indigenous African timber suckers and wildlings are sometimes also used trees with similar wood, e. g. Britondophrogmo for propagation. Smallleafy cuttings treated and Khoyo species, may be a better option in with growth hormone root fairly easily, espe- many regions. clany when taken from 2-year-old plants Major references CAB International, 2005; In plantations close spacing of young trees is Gintings at a1. , 1995; Katende, Birnie & recommended to prevent early branching. The Tengnds, 1995; Takahashi, 1978; World Agro- most commonly reported spacing is 2 in x 2 in, forestry Centre, undated but marrica 4-6 in x 4-6 in is commonly used Other references Burki11, 1997; Chilufya & Regular weeding is necessary during the first 2 Tengiias, 1996; CTFT, 1962c; Forestry Re- years because the seedlings are very sensitlve search Institute, 1981; Gelfand at a1. , 1985; to competition from herbs. Mahberley, Pannell & Sing, 1995; Malairajan Thinning operations should be done from the at a1. , 2007; Phiri, 1997; Sinoya, 2003; Styles & 4th year, and then every 5 years. Trees can be White, 1991. managed by copplcmg and pollarding. Authors R. H. M. J. Lemmens Leaf blight caused by Phytophthoro has been reported from forest nurseries in India. In

. 560 TIMBERSl

TRICHILIAGILGIANA Harms strongly branched; young branches short-hairy. Leaves alternate, imparipinnately compound Protologue Bot. Jahrb. Syst. 23: 161 (1896) with (2-)5-8 pairs of leaflets; stipules absent; Family Menaceae petiole 4-10 cm long, rachis (5-)10-26 cm long; Origin and geographic distribution Trichi- petiolules 4-10 min long; leaflets opposite, nar- Jin gilgiono occurs from southern Nigeria east rowly elliptical to narrowly ovate or narrowly to eastern DR Congo, and south to northern obovate, 5-28 cm x 1.5-8 cm, curieate to obtuse Angola (Cabinda) at base, acuminate at apex, glabrous but glan- Uses The wood is suitable for light construc- dular dotted, pinnately veined. Inflorescence tion, interior trim, vehicle bodies, furniture, an axillary panicle up to 20(-30) cm long, cabinet work, boxes, crates, toys, novelties, short-hairy; bracts elliptical to ovate, up to veneer, plywood, hardboard and particle board, 7(-10) mm long, usually persistent. Flowers as wellas pulpwood for paper production unisexual, male and female flowers very SImi- In Congo the bark is considered to have anal- Iar in appearance, regular, 5-merous, creamy gesic and stimulant properties and is used in brown to pink, fragrant; pedice1 3-5(-8) mm traditional medicine to treat abdominal, chest long; receptacle cylindrical, I-3 mm long; calyx and fever pains, and as a tonic. In DR Congo cup-shaped, 2-3 min long, lobes 1.5-2 min sap of young leaves is applied to circumcision long; petals free, narrowly elliptical to nar- wounds and small quantities of pulverized rowly obovate, 7-10(-11.5) mm long; stamens leaves are added to drinking water to treat 4.5-6.5 mm long, fused at base into a tube, respiratory problems in children hairy inside; ovary superior, pyramidal, c. 2 Properties The heartwood is greyish white min x 2-3 min, densely hairy, 3-celled, style to pinkish grey or pinkish brown and iridis- 2.5-4.5 mm long, hairy, stigma head-shaped; tinctly demarcated from the creamy white male flowers with rudimentary ovary, female sapwood. The grain Is often interlocked, tex- flowers with non-dehiscing arithers. Fruit a ture fine to moderately coarse. Quarter-sawn pear-shaped to fig-shaped capsule 3-3.5 cm in wood is lustrous and has darker stripes and diameter, with 0.5-I cm long stipe, dehiscent, Irregular bands up to 6-seeded. Seeds c. 20 min x 12 mm, on a The wood is medium-weight, with a density of long funicle, seed coat partly fleshy and or- 570-650 kg/ina at 12% moisture content. It air ange-red, remaining part glossy dark brown dries moderately well. The wood is not stable Seedling with epigeal germination; hypocotylc In service At 12% moisture content, the 4 cm long, epicotyl c. 4.5 cm long; cotyledons modulus of rupture of wood from Mayombe sessile, thick and fleshy (Congo) is 96 N/min', modulus of elasticity The flowers are pollinated by insects such as 9200 N/mm', compression parallel to grain 45 bees. The seeds are eaten by birds, which may N/mm', cleavage 13 N/mm' and Chalais- play an important role in dispersal Meudon hardness 2.7. Trichilio comprises about 90 species, most of The wood is difficult to saw because of the high them in tropical America. In continental Africa silica content, which causes blunting of saw 18 species occur, in Madagascar 6 teeth and cutting edges. It planes satisfacto- The area of distribution of Trichilio retuso rily, but quarter-sawn surfaces may be rough Onv. overlaps largely with that of Trichilio because of the presence of interlocked grain. gilgiono, but it is slightly larger, extending to The nailing properties are good, gluing proper- southern Sudan and south-western Ethiopia in ties moderate. The wood is not durable; it is the north, and southern DR Congo in the susceptible to fungal, termite, dry-wood borer south. Trichilio retuso is a shrub to small tree and marine borer attacks. The heartwood is up to 15 in tall, characterized by its broadly resistant to Impregnation by preservatives, but notched leaflets. In Sudan the wood is used for the sapwood Is moderately permeable. The pestles and mortars, whereas the fleshy seed wood dust may cause irritation to the respira- coat is reportedIy edible. In Congo bark scrap- tory tracts and skin in wood workers Ings are applied to swellings Botany Evergreen, smallto medium-sized tree Trichiliu rubescens Onv. has a similar distribu- up to 30 in tall; hole branchless for up to 18 in, tion, but extends to western Uganda and west- straight, cylindrical or fluted, up to 100 cm in ern Tanzania. It is also a shrub or small tree, diameter; bark surface smooth, greyish brown, differing from other African fitchi!to species in peeling off in small plates, inner bark pale its comparatively smallflowers with distinct pink, exuding some latex; crown spreading, disk and obovoid to globose fruits I-2 cm in

. TRiCHiLIA 561 diameter. The wood is used for poles, small east to the Central African Republic and DR implements and tool handles. It is also used as Congo, and south to northern Angola fuelwood and for making charcoal. Bark decoc- (Cabinda) tions are used in traditional medicine to treat Uses The wood is used in house building, bronchitis, stomach-ache, diarrhoea, dizziness, especially for piles. It is locally favoured for insanity and sexual impotence. They are also wood carving, e. g. for making masks and ca- administered as an enema to treat constipation noes. It is suitable for light flooring, Joinery, and abdominal complaints. Root decoctions are interior trim, ship building, vehicle bodies, applied as an enema to treat colic, and as furniture, cabinet work, boxes, crates, toys, arithelmintic and abortifacient. Young leaves novelties, veneer, plywood, hardboard and par- are taken to treat gonorrhoea. The bark is used ticle board, as well as pulpwood for paper pro- for making arrow poison. The fleshy seed coat duction. It is also used as firewood and for is eaten in times of food scarcity. Antiparasitic charcoal production. 11monoids were isolated from Trichilio rubes- Trichilio monodelpho is an Important medici- certs leaves, and leaf extracts showed antima- nal plant, and particularly Its bark is coin- Ianal activity. monly used in traditional medicine. A bark Ecology Trichilio gi!giono is a common tree decoction or the pulped bark is applied exter- in lowland evergreen and semi-deciduous sec- nany to wounds, sores, skin affections includ- ondary forest, often in the understorey, up to ing yaws, lumbago and oedema. A bark decoc- 950 in altitude tion is drunk to sooth cough, as an analgesic Management After felling, logs should be and arithelmintic, and to treat gonorrhoea and removed rapidly from the forest to dry, to pre- syphilis, whereas small amounts of pulped vent fungal and insect attacks, or they should bark are eaten or applied as an enema to treat be treated with preservatives gastro-intestinal complaints. Bark decoctions Genetic resources and breeding Trichilio serve as an aphrodisiac, ecbolic and abortifa- gilgiono does riot suffer from genetic erosion cient. A leaf decoction is taken to treat heart because it is fairly widespread and locally complaints, and pounded leaves to treat gonor- common in secondary forest rhoea and lumbago. The roots are an Ingredi- Prospects The wood of Trichilio gilgiono Is ent in preparations to treat dysentery, and are not particularly attractive and it Is abrasive, considered aphrodisiac. while the often fluted boles hamper Its applica- The seed o111s occasionally used in cooking. hinty in the peeling industry. Therefore, the The reddish brown dye present in the bark has prospects as a commercial timber tree seem to been used for dyeing cloth and hides. fitchiliu be poor, and filchi!to gilgiono may remain an monode!pho is useful for soil protection and undesirable tree in exploited forest, as it often soilimprovement has been, like other Tri, hilto species Properties The heartwood is pale plnkish Major references Bolza & Keating, 1972; brown to reddish brown and more or less dis- Burki11, 1997; de Wilde, 1986; Fouarge & tinctly demarcated from the whitish brown or G6rard, 1964 greyish brown sapwood. The grain Is usually Other references Keay, 1958b; Keay, 1989; straight, texture fine to moderately coarse. The thief at a1. , 2006; Krief at a1. , 2004; Lovett at wood contains latex cells and has a cedar-like a1. , 2006; Neuwinger, 2000; Staner & GIIbert, odour 1958; Terashima & Ichikawa, 2003 The wood is moderately lightweight, with a Authors R. H. M. J. Lemmens density of 510-580 kg/ms at 12% moisture content. The rates of shrinkage during drying are moderate. The wood is moderately stable in TRlcHiLIAMONADELPHA (Thonn. ) J. J. de service. It is soft. At 12% moisture content, the Wilde modulus of rupture is 63-110 N/mm2, modulus of elasticity 9200-9800 N/min2, compression Protologue Acta Bot. Neer1. 14: 455 (1966). parallel to grain 40-45 N/mm2, Janka side Family Menaceae hardness 2820 N, Janka end hardness 4270 N Chromosome number2n= 50 and Chalais-Mendon hardness 2.7 Synonyms ritehilio heudelotiiPlanch. ex 011v The wood is rather difficult to saw and work (1868) because of quickly gumming up and blunting of Origin and geographic distribution Trichi- sawteeth and cutting edges. It planes and fin- lid monodelpho occurs from Guinea Bissau ishes satisfactorily, producing nice hacksawn 562 TIMBERSl

and quarter-sawn surfaces. The nailing and ling with epigeal germination; hypocoty1 1.5-4 gluing properties are good. The wood is moder- cm long, epicoty1 2-3.5 cm long; cotyledons ately durable; it is susceptible to blue stain, sessile, thick and fleshy, green dry-wood borer and marine borer attacks, Flowering often coincides with the develop- whereas the liability to termite attack is re- merit of terminal buds and formation of new corded as variable. The heartwood is resistant shoots. In Nigeria Trichilio monodelpho now- to impregnation with preservatives, but the ers in the dry season. The flowers are polli- sapwood is permeable. The sawdust may cause nated by insects such as bees. In Guinea the irritation to the respiratory tracts in wood fruits ripen in December, 'anuary. The seeds workers are eaten by birds, which may play an impor- The presence of alkaloids and tannins has been tant role in their dispersal. demonstrated for the bark. Limonoids have Trichilio comprises about 90 species, most of been isolated from bark and roots. Bark ex- them in tropical America. In continental Africa tracts showed antiplasmodial activity against 18 species occur, in Madagascar 6. chloroquine- and pyrimethamine-resistant Trichilio dialonis Achev. , a shrub or small PIOsmodium 101cipurum strains (IC5o 3.6 tree up to 15 in tall occurring from Guinea to 11g/inI). The leaves contain tannins and flavon- C6te d'Ivoire, closely resembles Trichi!to olds. Leaf extracts showed activity against sev- monode!pho, but differs in usually fewer, gla- eral bacteria and fungi, including significant brous leaflets and smaller fruits. A reddish dye activity against the plant-pathogenic fungus can be prepared from the bark of Trichilio djo- Pusorium orysporum 10nis, and probably Its wood is occasionally Botany Evergreen, smallto medium-sized tree used for similar purposes as that of Trichilio up to 20 in tall; bole straight and cylindrical, monode!pho. often low-branching, up to 40(-60) cm in di- Trichi!io omithothero J. J. de Wilde, which is a ameter, without buttresses; bark surface small tree up to 15(-20) in tall occurring from smooth, pale grey to greenish brown or dark Sierra Leone to Ghana, is also close to Trichilio brown, inner bark pale brown to pink; crown monode!pho and there is much confusion in the spreading, open; young branches short-hairy. literature. Undoubtedly the wood is used for Leaves alternate, Imparipinnately compound similar purposes, and the bark in traditional with 3-6(-7) pairs of leaflets; stipules absent; medicine. Trichiliu omithotheru is character- petiole 4-13 cm long, rachis (4-)8-20(-23) cm ized by generally larger and more hairy leaves long; petiolules 2-7(-11) min long; leaflets op- in comparison with Trichiliu monodelpho, and POSite, ovate to obovate, 4-26 cm X 1.5-9 cm, by its usually 2-celled ovary. fitchtlio orni- curieate to obtuse at base, acuminate at apex, thothero is classified as vulnerable in the hairy below when young but glabrescent, pin- IUCN Red List. nately veined. Inflorescence an axillary panicle Trichilio tessmonnii Harms (synonyms: 77i- up to 12(-21) cm long, short-hairy; bracts ovate chino longto A. Chev. , Trichilto mildbroedii or triangular, up to 5 min long, caducous. Harms) has nearly the same distribution area Flowers unisexual, male and female flowers as Trichilio monode!pho, butts in most regions very similar in appearance, regular, 5-meTous, less common. It Is a medium-sized tree up to 30 greenish yellow or greenish white, fragrant; in tall, with straight, cylindrical bole up to 80 pedicel up to 2 min long; receptacle cylindrical, cm in diameter, and Is most easily recognized I-2.5 mm long; calyx cup-shaped, 1.5-2.5 min by its twigs having reddish brown, flaking long, lobes I-1.5 mm long; petals free, nar- bark. It has been stated that the wood is used rowly oblong, 7-10 mm long; stamens 5.5-7.5 in house building and that it is resistant to min long, basal half fused into a tube, hairy termites. The bark is used to treat stomach- inside; ovary superior, globose, 2-3 mm in di- ache and as a purgative. In DR Congo cooked ameter, densely hairy, (2.3(-4)-celled, style fruits are eaten. In Nigeria the seeds are used 3-5 min long, hairy, stigma head-shaped; male for rattles and tambourines flowers with rudimentary ovary, female flowers In the past Trichilio weIruitschii CDC. has with non-dehiscing arithers. Fruit an obovoid to been much confused with Trichtlio monodel- globose capsule 1.5-2.5 cm in diameter, pho, but it differs in its 2-celled ovary and (2-)3(-4)-lobed, with short stipe, dehiscent, up fruit. It is a small to medium-sized tree up to to 6-seeded. Seeds 15-17 min x 8-11 min, on a 30 in tall with a cylindrical hole up to 45(-60) long funicle, seed coat partly fleshy and or- cm in diameter, which occurs from Nigeria to ange-red, remaining part glossy blackish. Seed- eastern DR Congo and northern Angola. The

. TRIGHiLiA 563 wood is probably used for similar purposes as at al. (Editors), 1988; Atindehou at a1. , 2004; that of litchilio monode!pho. In DR Congo a Fouarge & G6rard, 1964; Hawthorne & Jong- bark decoction is administered as an enema to kind, 2006; Holingren at a1. , 2004; Hubert, un- treat haemorrhoids and other abdominal disor- dated; Neuwinger, 2000; Of on, 1999; Staner & ders, and as an abortifacient, whereas pounded GIIbert, 1958. young leaves are applied to syphilitic sores. Authors R. H. M. J. Lemmens Trichilio gilletii De Wild. occurs in approximately the same region and closely resembles Trtchiliu weiwitschii, from which it differs in TRiCHiLLAPRiEUREANA A. Juss fewer and glabrous leaflets. Its wood is probably used similarly, whereas its bark is used to Protologue Bull. Sd. Nat. Geol. 23: 238 (1830) treat fever and as a purgative, and its seed oil Family Menaceae as an emetic. Chromosome number2n=50 Ecology fitchi!io monodelphu is a common Vernacular names Monkey apple (En). Mti- understorey tree in lowland evergreen and mail(Sw) semi-deciduous secondary forest, up to 650 in Origin and geographic distribution Trichi- altitude, also occurring in forest edges, some- lid prteureono is widespread from Senegal east times in deciduous forest, butthen along rivers to south-western Ethiopia, Uganda and west- and in other moist localities. ern Tanzania, and south to northern Angola Management The 1000-seed weightis about (Cabinda) and Zambia. 400 g. When sown immediately after harvest, Uses The wood is used in Ethiopia for the seeds germinate within 8-15 days, with a high construction of local houses. In Tanzania It Is germination percentage. Young seedlings re- used for tool handles and spoons. The wood is quire shade. In Guinea Trichilio monode!pho suitable for heavy and light construction, rail- was planted in full sun, in forests paths and way sleepers, heavy and light flooring, joinery, under cover in secondary forest. The best re- interior trim, furniture, cabinet work, sporting suits were obtained under cover, with a mortal- goods, toys, novelties, veneer, plywood, hard- ity of about 40% and a tree height of over 3 in board and particle board. It is used as firewood after 6 years, whereas in unshaded conditions and for charcoal production; It burns slowly mortality was over 70% and the trees were only with great heat. I in tall In West Africa the bark is used to treat vene- In Ghana stem bark is harvested throughout real diseases, fever, cough, constipation, poi- the year, in pieces up to 5 cm x 2.5 cm, to be soning and ascites, and as an aphrodisiac. In used for medicinal purposes, mainly for the the Central African Republic a bark decoction treatment of nausea and stomach-ache. The is applied to treat pain caused by lumbago and bark is cleaned, dried in the sun and stored at rheumatism. Powder madefromburned bark is a moisture content below 9%. Packages of 40 applied to scarification wounds. Leaves, bark kg are transported and roots are applied against arthrltis. A leaf Genetic resources and breeding Trichilio decoction is drunk against anaemia and is ap- monodelpho does not suffer from genetic ero- plied as a bath against syphilis, whereas pul- SIon because it is widespread and locally coin- verized leaves are taken to treat stomach mori in secondary forest. Locally, large-scale spasms A decoction of various plant parts Is harvesting the bark may severely reduce popu- applied as a wash to treatleprosy and wounds. Iations A decoction of leafy twigs Is taken to treat Prospects The small size of the hole of Tri- bronchitis and oedema. Twigs are used as chilio monodelpho limits Its importance on the chew-sticks. The root and pounded leaves are international timber market, although it is used in preparations to treat gonorrhoea, root locally important in house building, mainly as and bark are administered as an enema to piles. Although its bark is a common ingredient treat piles, and pulverized roots are taken of traditional medicinal preparations, little against ascariasis and as a purgatlve. The seed research has been done on lts phytochemistry is one of the ingredients of a preparation to and pharmacologicalactivity treat gottre. The fleshy seed coat is edible. In Major references Aubr6ville, 1959a; Bolza Gabon Trichilioprieureono has been used as a & Keating, 1972; Burki11, 1997; de Wilde, 1986; shade tree and support for van111a plants. Takahashi, 1978. Properties The heartwood is pale pinkish Other references Add0, 1998; Adjanohoun brown to reddish brown and distinctly demar- 564 TIMBERSl

cated from the creamy white to pale yellow epigeal germination; hypocotyl c. 4 cm long, sapwood. The grain Is wavy or straight, texture epicoty1 2.5-3 cm long; cotyledons sessile, thick fine and fleshy, green The wood is moderately heavy, with a density In Sierra Leone Trichi!toprieureono flowers in of about 750 kg/in3 at 12% moisture content. It January-March, and fruiting is from March air dries moderately well to with difficulty; the onwards. The flowers are pollinated by insects rates of shrinkage are moderately high. The such as bees. The fleshy seed coat has been wood is difficult to saw because of the presence recorded in Gabon as an important food for of silica, which causes blunting of saw teeth monkeys and birds such as hornbills and tura- and cutting edges. It planes satisfactorily, giv- cos early in the dry season Ing nice quarter-sawn surfaces. It polishes Trichilio comprises about 90 species, most of well. The nailing properties are good. The wood them in tropical America. In continental Africa Is moderately durable; it is susceptible to 18 species occur, in Madagascar 6 Lyetus attack. The heartwood is resistant to fitchi!to prteureonu has an isolated position impregnation by preservatives, but the sap- within the genus in Africa and has been placed wood is moderately permeable. The wood dust in the section MOSchoxylum. It is variable and may cause irritation to the respiratory tracts in 3 subspecies have been distinguished: subsp wood workers prteureono (synonym: fitchilio servegolensis Botany Evergreen shrub orsmallto medium- CDC. ) occurring from Senegal to Nigeria, and sized tree up to 30(-40) in tall; hole branchless characterized by usually 3-celled ovary, gla- for up to 21 in but usually much shorter, often brous style and lobed stigma; subsp uermoe- crooked or sinuous, usually conspicuously senti J. J. de Wilde occurring from C6te d'Ivoire fluted, up to 100 cm in diameter; bark surface to Uganda and Angola, and characterized by shallowly fissured, greyish brown, peeling off usually incompletely 2-celled ovary, slightly in thin flakes or rectangular strips, inner bark hairy style but glabrous ovary, and head- pale yellow or pink; crown hemispherical, shaped stigma; and subsp. orientalis J. J. de dense; young branches glabrous. Leaves alter- Wilde occurring in southern DR Congo, nate, imparipinnately compound with Uganda, western Tanzania and northern Zam- (I-)2-4(-5) pairs of leaflets; stipules absent; bia, and characterized by usually incompletely petiole (1.5-)3-10 cm long, rachis 2-celled ovary, slightly hairy style and ovary, (I-)4-15(-20) cm long; petiolules 2-10(-20) and head-shaped stigma min long; leaflets opposite, elliptical to ovate or Ecology Trichilio prtet, redno occurs in low- obovate, 5-25 cm x 2-10 cm, curieate at base, land forest and riverme forest up to 1300 in acuminate at apex, glabrous, pinnately veined altitude, up to 1500 in in Zambia, often as an Inflorescence an axillary panicle up to 10(-13) understorey tree. In West Africa it prefers cm long, short-hairy; bracts ovate to triangular, drier forest types, with subsp. prteureono usu- up to 2.5 min long, caducous. Flowers unisex- ally found in savanna woodland and forest- ual, male and female flowers very similar in savanna mosaic, and subsp. uermoesenii in the appearance, regular, 5-merous, greenish white, rainforest region. In Uganda subsp. Dermoe- fragrant; pedicel up to 2 min long; receptacle sen, ! occurs in rainforest in higher-rainfall cylindrical, up to 1.5 mm long; calyx cup- areas in the western part of the country, subsp shaped, I-2.5 min long, lobes 0.5-2 min long; orientalis in savanna woodland and forest- petals free, narrowly obovate to narrowly ob- savanna mosaic in the northern and eastern long, 4.5-8 min long; stamens 3-6 mm long, parts of the country fused completely into a tube, hairy inside; Management The 1000-seed weightis about ovary superior, ovoid to globose, I-2 mm in 330 g. Seeds germinate $15 days after sowing diameter, glabrous or slightly hairy, 2-3-celled, Logs split easily and therefore some care is style I-4 mm long, glabrous or slightly hairy, needed during felling operations stigma head-shaped or distinctly lobed; male Genetic resources and breeding Trichilio flowers with rudimentary ovary, female flowers prteureono does not suffer from genetic erosion with non-dehiscing arithers. Fruit an ovoid to because it is widespread and locally common, globose capsule 1.5-2.5 cm in diameter, often also in secondary forest. pink when ripe, dehiscent, up to 6-seeded Prospects The often sinuous and fluted boles Seeds 10-17 min x 7-12 mm, on a long funicle, of Trichiliopriet, redno hamper its applicability seed coat partly fleshy and orange-red, remain- in the peeling industry, and the presence of ing part glossy dark brown. Seedling with silica, which makes sawing difficult, and easy

. TRIPLOCHITON 565 splitting of the wood are other drawbacks joinery, panelling, moulding, furniture, boxes Therefore, the prospects as a commercialtim- and crates, sculptures, matches, pencils, peeled her tree seem to be poor, and Trichilioprieure- and sliced veneer for interior and exterior parts ono may remain an undesirable tree in ex- of plywood, fibre and particle boards, and PIOited forest, as it has often been, like other blockboard. It is of great importance for house Trichilio species building, for beams, posts and planks, and Is Major references Bolza & Keating, 1972; also used for roof shingles. The wood from the BurkiU, 1997; de Wilde, 1986; Styles & White, buttresses is used to make doors, platters, 1991; White & Abernethy, 1997. bowls and sandals, and the hole is used for Other references Adjanohoun at a1. , 1989; dugout canoes. The wood pulp can be used to Mobgninou, van der Burg & van der Maesen, produce paper of moderate quality 2006; de in Mensbruge, 1966; Hawthorne, The leaves are prepared as a cooked vegetable 1995; Hawthorne & Jongkind, 2006; Keay, or sauce in traditional cuisine in C6te d'TVoire 1989; Lovett at a1. , 2006; Motte, 1980; Neu- and Benin. The bark is used to cover the roof winger, 2000; Raponda-Walker & Sillans, 1961. and walls of huts. It is also applied in tradi- Authors R. H. M. J. Lemmens tional medicine to treat oedemas and as an anodyne. Triplochiton setoro, cylon is a food plant of the silkworm Ariaphe ue, Iota, the Iar- TRIPLOCHITONSCLEROXYl, ON K. Schum vae of which are a good source of protein and commonly eaten. Sawdust is used for the pro- Protologue Bot. Jabrb. 28: 331 (1900) duction of edible fungi (Pleurotus spp. ). The Family Sterculiaceae (APG: Malvaceae) trees are often preserved in cocoa plantations Chromosome number2n=40 to serve as shade trees Synonyms Triplochiton itigericum Sprague Production and international trade fliplo- (1909) chiton seieroxy!on used to be the major timber Vernacular names African whitewood, Mri- tree of West and Central Africa. In 1959 Ghana can maple, ayous, obeche, wawa (En). Ayous, exported 650,000 ing of logs and 30,000 ina of obeche, samba (Fr) sawn timber, while the export of logs from Ni- Origin and geographic distribution Triplo- gena was 350,000 in3, from Cote d'Ivoire chiton scleroxylon is widely distributed in the 91,000 ing and from Gamero0n 1750 in3. In West and Central African forest zone from 1973 Cote d'Ivoire exported over I million ms of Guinea east to the Central African Republic, logs and Ghana, Nigeria and Garnero0n to- and south to Gabon and DR Congo. It is coin- gether 400,000 ing of logs and 40,000 in3 of monly planted in its natural area of distribu- sawn timber. In 1983 export from 06te d'Ivoire tion (e. g. in C6te d'TVoire, Ghana and Nigeria), decreased to 230,000 ms. reflecting the loss in and occasionally elsewhere, e. g. in the Solomon forest area. Currently, it is economicalIy the Islands most important timber species of Ghana and Uses The wood is widely used for Interior Cameroon, making up about 70% of the volume of timber products exported from Ghana and 35% from Cameroon. In 1996 the volume of fliplochiton seieroxylon timber ('ayous') exported from Cameroon (mainly as logs) was estimated at 700,000 ina and in 1998 475,000 . ing. In 2001 the export of logs and sawn timber

. from Garnero0n amounted to 29,000 ing and 11,000 in3 respectively, according to ITTO statistics, with a mean price of Us$ 931m3 for logs .^ and Us$ 4351m3 for sawn timber. Also in 2001, the export of sawn flip!ochiton seiero, cy!on \.. timber from C6te d'TVoire ('samba') was 109,000 in3, with a mean price of Us$ 2201m3, ., and 2000 ms veneer was exported, with a mean

o price of Us$ 2871m3; the export of sawn timber from Ghana ('wawa') was 114,000 ina in 2001, with a mean price of Us$ 2461m3, and 80,000 Trip!ochiton seieroxylon - wild ing in 2002. In 2003 export from Cameroon was 566 TIMBERSl

86,000 in3 of logs, mainly to Italy and China, orylon and 282,000 ms of sawn timber, mainly to Italy Fresh leaves have a moisture content of 74.7%, and Spain and they contain per 100 g dry matter: energy Properties The heartwood is whitish to pale 668 kJ (160 kcal), protein 29.2 g, fat 2.2 g, yellow, Indistinctly demarcated from the sap- starch 3.0 g, sugar 2.3 g, fibre 51.0 g, Ca 1/14 wood, which is up to 15 cm thick. The grain is ing, Mg 551 ing, Fe 9.2 ing, p-carotene 16.5 ing, usually interlocked, sometimes straight, tex- riboflavin 0.78 ing and ascorbic acid 165 ing ture moderately coarse. The wood has a ribbon- (Herzog, Farah & Amado, 1993). The mucilage like aspect on quarter-sawn faces, and is jus- content of the leaves per 100 g was 4.2 g, coin- trous. Fresh wood has an unpleasant smell, pared to 9.1 g in the fruits of common okra which disappears upon drying (Abelmoschus esculentus (L. ) Moench) The wood of Triplochiton seieroxy!on is light- Adulterations and substitutes The wood weight, the density is 320-440(-490) kg/in3 at of Triplochiton seieroxy!on can be replaced by 12% moisture content. The shrinkage rates are other lightweight and whitish wood such as moderately low, from green to oven dry 2.5- that of AIStonio boonei De Wild. , Antioris tort- 4.1% radial and 4.2-6.6% tangential. The tim- curio Lesch. and Getbopentondro (L. ) Gaertn her dries easily and rapidly, with only a slight Description Large deciduous tree up to 50 risk of distortion and checking. The use of large in tall; bole straight, often angular and spacer sticks is recommended during air drying strongly ridged, branchless for up to 30 in, up to allow good air circulation. Once dry, the to 150(-210) cm in diameter, with low to very wood is stable in service high (up to 8 in) buttresses; bark 7-30 mm At 12% moisture content, the modulus of rup- thick, grey to yellowish brown, smooth when ture is 52-110 N/mm2 modulus of elasticity young, becoming scaly with age, often with 4800-9200 N/min2, compression parallel to vertical lines of lenticels; crown dense and grain 24-43 N/min2, shear 3-8 N/mm2 cleav- rounded, with heavy, not far-spreading age 5-15 N/min, Janka side hardness 1910- branches. Leaves alternate, simple; stipules 2100 N and Janka endhardness 3070-3600 N linear, 2-4 cm long, early caducous and leaving The wood works easily with hand and machine annular scars; petiole (1.5-)3-10 cm long; tools; ordinary saw teeth and cutting tools can blade palmately 5-7-10bed, up to 20 cm long be used. However, sharp edges are needed for a smooth finish, and the wood tends to tear in $\>:;. , mortising. A cutting angle of 15' is recoin- 3 \.;-,'.\\;\'

* --.~;.~* mended when planing to prevent tearing. The ;,/ peeling and slicing properties are good. The . .. wood stains and polishes well. Filling is recommended to obtain a good finish. The nailing and screwing properties are rather poor; split- , , ting may occur. Gluing does not cause prob- Iems, although it must be done carefully as the wood is absorbent. The carving characteristics are good The wood is not durable, being liable to fungal attack (e. g. blue stain), and susceptible to termites, powder-post beetles and dry-wood bor- t ers. It should not be used in contact with the ... *. ,. ' .,' I A ground or exposed to the weather. The heart- :.

, 2 wood is somewhat resistant to preservative ^. treatment. In an autoclave, the wood absorbs \.. I: . 140 Urns of preservative \-. I The sawdust may cause occupational allergic . .., contact dermatitis, allergic rhinitis and * .. 11 asthma ~<3f, *.,,".' ' ' ' ~.,^ ", The pulping properties of the wood are moderate. Unbleached and bleached kraft pulps of 7,4010chiton seieroxylon - I, hose of 601e, 2, flow- good yield and quality were made from a mix- ering twig, . 3, fruit ture of wood chips including Triplochiton scler- Redrown und odopted by Ishoh Syomsudin

. TRIPLOCHiTON 567 and wide, cordate at base, lobes broadly ovate ginal cells; 110: sheath cells present; 1/1: tile or triangular, obtuse to acute at apex, brown cells present; 1/5: 4-12 rays per mm Stoned stellate hairy when young but soon becoming structure: 1/9:low rays stoned, high rays non- glabrous. Inflorescence an axillary or terminal stoned; 120: axial parenchyma and/or vessel panicle up to 10 cm long, densely hairy; bracts elements stoned; 121: fibres stoned. Mineral c. 0.5 cm long, early caducous. Flowers bisex- inclusions: 136: prismatic crystals present; 137: ual, regular, 5-merous; pedice1 3-4 mm long, prismatic crystals in upright and/or square ray articulate at base; sepals triangular, c. 7 mm cells; 141: prismatic crystals in non-chambered long, brown stellately hairy; petals broadly axial parenchyma cells obovate, c. I cm long, pinkish white but purple (N. P. Monel, P. 06tienne & E. A. Wheeler) at base, densely silky hairy; androgynophore c. Growth and development Under natural 3 mm long, hairy; stamens 30-40, connate at conditions seedlings may reach 15 in tall and base in pairs; ovary surrounded by 5 petal-like 15 cm in stem diameter after 4 years. Mean staininodes, consisting of 5 free carpels, styles annual diameter increment in the forest aver- connate. Fruit consisting of I-5 rhombic nuts c ages I cm, but in Nigeria mean annual diame- 2 cm x I cm with a large wing 4-6 cm x I-2 ter increments of up to 2.5 cm have been re- cm. Seedling with epigeal germination; hypo- corded and in the Central African Republic as cotyl c. 7 cm long, epicoty1 1.5-2 cm long; coty- low as 03 cm. In a 19-year-old plantation in ledons leafy, rounded, 2.5-3 cm in diameter, 3- Ghana with 600 stemsftia, the trees were on 5-veined from the base. average 21.8 in tall with a hole diameter of 27 Other botanical information TrLplochiton cm. In Nigeria half of the annual diameter In- comprises only 2 species. It resembles Mon- crement takes place from inId April to inid sonio, which differs in its entire leaves and July. spathaceous calyx. fliplochiton gumbesiocus The hole and branches of Triplochiton seieroxy- Milne-Redh. occurs in southern Africa 10n are all orthotropic and branching is rhyth- Anatomy Wood-anatomical description OAWA inic, corresponding to Rauh's model of tree ar- hardwood codes): chitecture. Normally, the trees do not flower Growth rings: (I: growth ring boundaries dis- until 15 years old. In Ghana and Nigeria flow- tinct); (2: growth ring boundaries indistinct or ering occurs during the long dry season be- absent). Vessels: 5: wood diffuse-porous; 13: tween November and March when the trees simple perforation plates; 22: intervessel pits have a leaness period. In Liberia trees flower alternate; 23: shape of alternate pits polygonal; in DecemberJanuary, when the leaves are 26: intervessel pits medium (7-10 F1m); 30: usually shed. It has been observed that flower vessel-ray pits with distinct borders; similar to initiation is associated with changing tempera- intervessel pits in size. and shape throughout ture. The fragrant flowers open late in the day the ray cell; 42: mean tangential diameter of and wither within 18 hours; they are pollinated vessellumina 100-200 pin; 43: mean tangen- by insects. Cross poUination is needed for the tialdiameter of vessellumina z 200 prn; 46: S 5 production of viable seed. The trees produce vessels per square minimetre;(47: 5-20 vessels fruits very irregularly, but mast production per square minimetre); (56: tyloses common). occurs with intervals of several years. Fruit Tracheids and fibres: 61: fibres with simple to development extends into the start of the rainy minutely bordered pits; 66: non-septate fibres season, and is frequently impaired by pests, present; 69: fibres thin- to thick-waned. Axial such as the fruit-boring weevilApion ghonoe, I- parenchyma: 77: axial parenchyma diffuse-in- sis and pathogens including the smut fungus aggregates; (86: axial parenchyma in narrow Mycosyrin, c sp. The fruits are dispersed by bands or lines up to three cells wide); (89: axial wind. In Liberia ripe fruits are present in parenchyma in marginal or in seemingly mar- January-March. ginal bands); 90: fusiform parenchyma cells; Ecology fliplochiton scleroxylon is charac- 91: two ceUs per parenchyma strand; 92: four tenstic for semi-deciduous forest, where it of- (3-4) cells per parenchyma strand. Rays: 98: ten grows gregariously, but it can sometimes larger rays commonly 4- to 10-senate; (99: jar- be found in clearings in dense evergreen forest ger rays commonly > 10-senate); 102: ray and in dry forest. In Nigeria It is almost exclu- height > I min; 106: body ray cells procumbent SIvely limited to moist forest areas at low and with one row of upright and/or square marginal medium altitudes. It occurs up to 900 in alti- cells; 107: body ray cells procumbent with tude in regions with an annual rainfallofup to mostly 2-4 rows of upright and/or square mar- 3000 min, butts most abundant at 200-400 in 568 TIMBERSl altitude and in areas with an annual rainfallof Planting outis usually done at the beginning of 1100-1800 min and 2 rainy seasons. It prefers the rainy season. Normally potted seedlings of more fertile, well-drained, rerruginous soils about 30 cm tall are planted in holes 40 cm with light or medium texture and acid to neu- wide, after cutting the base of the pot to re- tral pH. It does not tolerate waterlogging, and move deformed roots. In 2-3 weeks the stumps in general avoids swamps. It is a light-deman- develop several sprouts, of which one takes the ding pioneer species. Seedlings may be very lead. The survival rate of stumps of one year abundant in forest gaps of larger sizes, and the old (2.5-4 cm in diameter) is usually 50-60% tree Is characteristic of secondary forest They are liable to termite attack and to sup- Propagation and planting Most fruits col- pression by weeds. When planted at 3 in x 3 in Iected from the ground have been attacked by spacings, the canopy can close after 3 years insects. The fruits can be collected from the In Ghana planting under a taungya manage- trees when still green just before maturation. merit system has given good results. SIIvicul- One kg comprises about 3000 winged fruits tural practice in C6te d'Ivoire is directed to- Seeds start to germinate I-2 weeks after sow- wards maximum sunlight for tree crowns and ing, but the germination rate is often low limiting competition by early and regular clear- Fruits stored at -18'C fully retained their vi- ings. In general 50% of the trees are cut when ability for 18 months. However, at 25'C viabit- they have reached 6-7 in tall ity drops from about 80% to 15% after 6 Management Production of rimlochiton scle- months. Fruits stored at 8-15% moisture con- 70xylon timber Is usually in natural forest un- tent lost only a little viahinty after 12 months, der selective logging. In C6te d'Ivoire Triplochi- but soon thereafter viahinty declined sharply; ton setoroxylon has been planted since 1930 in when stored at 30% moisture content they lost mixed timber plantations, e. g. with Khoyo and their viahinty quickly. More than 50% of the Terminglid spp. From 1967 to 1995 about 3000 seeds may still germinate after being stored for ha was planted in C6te d'TVoire. Plantations 7.5 years in sealed containers at 4'C. Germina- have a cutting cycle of less than 40 years. In tion rate and speed increase when the seeds Nigeria Triplochiton seieroxylon is planted in are pre-treated by moistening between layers agroforestry systems with cocoa. The results of of damp cotton wool. Fruits with wings re- plantations in the Solomon Islands have been moved are shallowly buried in pots, which are rated as good, with well-formed, fast growing placed under shade. The seedlings are pricked and self-pruning trees out when first leaves appear. They are fragile Diseases and pests The roots are very sen- and sensitive to damping off. They develop a sitrve to fungal rot. In Nigeria the cricket taproot, which is often forked, with few lateral Gymnogry!jus Iucens, the grasshopper Zorioce- roots In the upper 15 cm of the soil. The taproot FMS Donegotus and the psyllid Dielidophlebio of a 2 in tansapling may be I in long. Seedlings sp. can cause serious damage to seedlings, and of I-3 years old with their tops intact but the the wood borers EUlophonotus obesus and root system severely cut back to permit plant- Trochyostus ghonoensis to adult trees. The Ing in 40 cm x 40 cm x 40 cm holes were suc- silkworm Anuphe Defterto is a defoliator of the cessful. However, it has also been observed trees. Beetles such as the bostrychid Apote that cutting the primary root for planting monochus bore holes up to I cm in diameter causes serious drawbacks; new roots form into the wood. slowly, which makes the plant liable to para- Harvesting Harvesting is done throughout sitic attacks. Single-node cuttings with one leaf the year. It involves felling and cross-cutting of were rooted successfully under mist; rooting the logs using power chain saws and skidding was Improved by a hormone dip and a high the log parts to a central log yard. The mini- temperature in the bed (30'0. Cuttings 10 cm mum diameter limit for cutting is 60 cm in long with 2-4 leaves taken from 2-month-old C6te d'Ivoire, 80 cm in Garnero0n and 90 cm in branches are also used. These are placed in a Ghana. Defects of logs include brittle heart, nursery under 40-60% shade under mist. It ring shakes, black holes and grub holes takes about 12 weeks to obtain a rooted and Yield In Ghana the total standing volume hardened plantlet. Marcotting is possible by was estimated at 3930 in31km2 in inventories in ringing a branch at an internode; results are 2001, and exploitable volume 1650 in31km2. The optlinal when the foliage is at maximum den- annual allowable cut has been fixed at 816 sity, between August and October, with up to ruskm2. In the 1960s, total standing volumes 50% success in 12-year-old trees above 60 cm diameter were estimated at 2350

. TURRAEANTHUS 569 in31km2 in Cameroon, 1410 in31km2 in Cote Other references Ashiru, 1988; Ashiru & d'TVoire, 1350 ms/kin2 in the Central African Mornodu, 1981; Attab, Bues & Sagor, 2005; Republic and 610 in31km2 in Congo BOSch, 2004; Dick at a1. , 2004; Germain & The final harvest in plantations in Cote Barnps, 1963; Hall & Swame, 1981; Hall6, d'Ivoire yields 200-250 mama of timber, of 1961; Hawthorne & Abu Juam, 1995; Inside- which 170-200 ina from the hole, with an an- Wood, undated; ITvine, 1961; Leakey, 1992; nualvolume increment of8-13 ms/ha Nketiah, Newton & Leakey, 1999; Onnude & Handling after harvest Logs should be re- Ogunsanw0, 2002; Takahashi, 1978; Wagner, moved from the forest or treated as soon as Atuahene & Cobbinah, 1991; Wilks & TSSemb6, possible after cutting because they are suscep- 2000; World Agroforestry Centre, undated. tmle to fungal and insect attacks. Logs of Tri- Sources of illustration Hall6, 1961; Voor- PIOchiton sc!eroxy!on float in water, and can hoeve, 1979; Wilks & Issemb6, 2000 therefore be transported by river. In Ghana Authors FF. BOSu & E. Krampah logs are usually transported to processing mills or log markets by means of articulated hauling rigs TURRAEANTHUSAFRICANUS(WeIw. exC. DC. ) Genetic resources As a widely distributed Fellegr. pioneer species TrLplochiton seieroxylon Is not easily liable to genetic erosion, but in many Protologue Notul. Syst. (Paris) 2: 16, 68 regions within its distribution area it is under (1911) pressure because of excessive exploitation. In Family Menaceae Ghana it is considered vulnerable; it is still Chromosome number2n= c. 280 common, but under profound pressure. In Libe- Synonyms 7147roeonthus DigiteiHutch. & Dal- ria stocks are limited. Gene banks and clonal tie1(1928). trials representing the whole geographical Vernacular names Avodire (En). Avodir6 range of Trip!DChito, , seieroxylon have been (ET) established at 5 sites in Nigeria. In C6te Origin and geographic distribution furroe- d'TVoire slightly over 100 superior trees have grithus o17iconus is widespread, from Sierra been selected from a stand of over 50,000 trees Leone east to western Uganda, and south to to serve as mother trees for the large-scale DR Congo and northern ingola. production of cuttings. Uses The wood (trade names: avodire, avodir6) Breeding Selection of superior germplasm is valued for high-quality furniture, cabinet using DNA techniques ls currently being PUT- making, decorative carpentry, moulding and sued in Ghana panelling, musical instruments, and for sliced Prospects There are good opportunities for veneer. It is suitable for light construction, highly productive clonal timber plantations of flooring, ship building, vehicle bodies, toys, Trip!DChito, ! seieroxylon. In tests, wood mate- novelties, boxes, crates, vats, turnery, hard- rialfrom plantation-grown trees was found not board, particle board and pulpwood for paper inferior to that from trees harvested in natural forest. The erratic seed production is a major drawback for the establishment of plantations, but methods of relatively cheap vegetatlve propagation are possible and will offer great . opportunities when superior germplasm becomes available. The high growth rates, allowing comparatively short cutting cycles, the generally good form of the holes, and the possi- hinty of planting in mixtures with other timber ,^

species make flip!ochiton seieroxylon even L. more promising Major references Burki11, 2000; Chudnoff, .. 1980; CTRAD Forestry Department, 2003; CTFT, 1976c; Herzog, Farah & Amado, 1993; o Pana & Louppe, 2002; Richter & Danwitz, 2000; SIepel, Poorter & Hawthorne, 2004; Ver- nay, 2005; Voorhoeve, 1979. Turrueonthus @1nconus- wild

. 570 TIMBERSl

production. It is also used as firewood and for The wood is not durable, being susceptible to charcoal production fungal, dry-wood borer, termite and marine The bark is used in traditional medicine. Bark borer attacks. The heartwood is resistant to extracts are taken or added to a bath to treat impregnation with preservatives, the sapwood cough, fever, headache, epilepsy, filariasis, and is more permeable. The sawdust is very Irri- as an abortifacient. Dried and pulverized bark tant and may even cause internal bleeding in with salt added is rubbed into scarifications wood workers; good ventilation is required against hernia. The bark is also used as fish Several diterpenoids and triterpenoids have poison; leaves are occasionally used for the been isolated from the bark and seeds. Stem same purpose. The seed oilis used as an aborti- bark extracts and some of the isolated diterpe- focient noids exhibited significant antimicrobial activi- Production and international trade The ties against the pathogenic fungus Cryptococ- wood of Turroeonthus of neonus is traded on cus neoformons and the bacterium Stophy!0- the international timber market, where it coccus guruus. Alkaloids have also been ISO- fetches high prices for high-quality furniture lated from the stem bark. Bark extracts and decorative joinery. However, since the showed some activity against the storage pest 1960s the volumes exported are very small beetles Collosobruchz, s maculotus and Sitophi- C6te d'TVoire exported about 5000 ms of logs ms zeomois. A labdane diterpenoid (methyl per year between 1960 and 1974. Ghana ex- 14.15-epoxylabda-8(17), 12E-dien-16-oate) ISO- ported 3000 ing of logs in 1998; but only 150 in3 lated from the bark exhibited in-vitro anti- In 2001 PIasmodial activity against a chloroquine- Properties The heartwood is creamy white resistant strain of Plusmodium foieiporum to pale yellow, darkening to golden yellow upon T}Ie ent-labdane (+)-12.15-cookylabda-8(17), 12.14- exposure to light; it Is indistinctly demarcated trien-16-yl acetate isolated from the seeds from the 5-6 cm wide sapwood. The grain is showed in-vitro cytotoxic effects on cancer cell straight to interlocked, texture fine and even lines. The wood Is distinctly lustrous, with a silky Adulterations and substitutes The wood shine. Quartersawn surfaces often have an of TMrroeonthus of neoni, s has some resem. attractive inottled figure. blance to the wood of AIStonio boonei De Wild. , The wood is medium weight, with a density of Antioris toxicorio Lesch. , Curiorittm schwein- 480-660 kg/ms at 12% moisture content. With furthii Engl. , Porkio bicolor Achev. , Pterygoto some care, it air dries fairly easily and rapidly, inocrocurpu K. Schum. and Terminolio iuoreii- but there is a serious risk of distortion and sis A. Chev. , and serves in Ghana as a substi- slight risk of checking. The shrinking rates are tute for them. In Europe it is used as a substi- medium, from green to oven dry 3.4-4.8% ra- tute for European beech (Fagus), European oak dial and 5.3-7.9% tangential. Once dry, the (Quercus) and sycamore (Acer) wood Is very stable in service. Description Evergreen medium-sized to fairly At 12% moisture content, the modulus of rup- large tree up to 35(-45) in tall; hole branchless ture is (69-)83-166 N/min2, modulus of elastic- for up to 15(-30) in, up to 100(-120) cm in di- Ity 8300-12,100 N/min2, compression parallel ameter, fluted or with short buttresses at base, to grain 36-61 N/min2, shear 9-16 N/min2 sometimes extending in large surface roots; cleavage 13-20 N/min, Janka side hardness bark surface smooth to scaly, sometimes shal- 4800 N and Janka end hardness 6400 N lowly fissured and with vertical rows of lenti- The wood saws and works well with ordinary cels, grey to pale brown, Inner bark pale yellow equipment and with only slight dulling effects to pale brown, with orange speckles and cedar- on saw teeth and cutting edges. Tearing may like smell; crown Irregular, spreading, dark occur in planing because of the presence of green, with ascending branches; twigs densely interlocked grain; a cutting angle of 15-20' is rusty short-hairy, glabrescent. Leaves alter- then recommended. The wood can be polished nate, clustered in lax groups at ends of to an excellent finish. Splitting is common dur- branches, panpinnately or imparipinnately ing nailing and screwing, and pre-boring is compound with (5.8-36 leaflets; stipules ab- recommended. The gluing, painting and var- sent; petiole 5-17 cm long, swollen and slightly nishing properties are allgood, butthe bending winged at base, rachis up to 60 cm long; petio- properties are poor. Sliced veneer of excellent Iules 5-10 mm long; leaflets alternate to OPPo- quality can be produced, but the logs are often site, oblong-elliptical to oblong-lanceolate, 6-29 too Irregular to produce good results in peeling cm x 2-6G8) cm, curieate to rounded at base,

. TURRAEANTHUS 571

PIe, with long-acuminate apex I Other botanical information furroeonthus ..~ I comprises 2-3 species and is confined to tropi- I, cal Africa. It seems closely related to Guoreo, which differs in usually having the staminal tube free from the corolla and in wood- anatomical characteristics.

I. Anatomy Wood-anatomical description (IAWA hardwood codes): 3 it Growth rings: 2: growth ring boundaries iridis- I~' tinct or absent. Vessels: 5: wood diffuse-porous; 13: simple perforation plates; 22: intervessel ^ pits alternate; 23?: shape of alternate pits po- Iygona1; 24: intervessel pits minute (S 4 pin); \!* 2 30: vessel-ray pits with distinct borders; SImi- * Iar to intervessel pits in size and shape ." throughout the ray cell; 42: mean tangential diameter of vessellumina 100-200 pin; 47: 5-

., 20 vessels per square minimetre; 58: gums and , other deposits in heartwood vessels. Tracheids and fibres: 61: fibres with simple to minutely . ..,

^ bordered pits; (65: septate fibres present); 66:

. " non-septate fibres present; 68: fibres very thin- .. walled; 69: fibres thin- to thick-walled. Axial 4 -, ;*;;;',, , - .*-.*...... parenchyma: 78: axial parenchyma scanty paratrachea1; 79: axial parenchyma vasicen- furroeonthus of neonus - I, base of 601e, ' 2, twig tric; 92: four (3-4) ceUs per parenchyma with led/; 3, brunch with inflorescences, 4, brunch strand; 93: eight (5-8) cells per parenchyma with inIructescence strand. Rays: 97: ray width I-3 cells; 104: all Redrown und adopted by AChmod Sotiri Nur- ray cells procumbent; 106: body ray cells pro- hornon cumbent with one row of upright and/or square marginal cells; 1/5: 4-12 rays per min. Mineral shortly acuminate at apex, tip with folded inclusions: 136: prismatic crystals present; 137: margins, leathery, initially short-hairy below prismatic crystals in upright and/or square ray but soon glabrescent, minutely scaly, pinnately cells; 138: prismatic crystals in procumbent ray veined with 10-30 pairs of lateral veins. Inflo- ceUs; (142: prismatic crystals in chambered rescence an axillary panicle up to 70 cm long, axial parenchyma cells) often on older branches, densely rusty hairy (E. Uetimane, H. Beeckman & P. E. Gasson) Flowers functionally unisexual, regular, 5- Growth and development Youngseedlings merous; pedicel c. 2 min long, jointed at base; require shade, but saplings need more light for calyx saucer-shaped, I-2 mm long, densely proper development, preferring small gaps in hairy; corolla tube 1.5-2 cm long, lobes 0.5-I the forest. Too much light, however, often re- cm long, hairy outside, creamy white to SUIts in low branching. In tests in Guinea, all brownish yellow; male flowers with stamens seedlings planted in full sun had died after 2 fused into a cup-shaped tube united with the years, whereas more than 80% of the seedlings corolla, with 10 included arithers, ovary not planted in forest understorey had survived functional; female flowers with superior, coin- after 3 years. Growth was slow, however, with cal ovary, 4-5-ceUed, graduaUy passing Into a mean height of 150 cm after 6 years' the style, stigma disk-shaped, stamens not The hole is often irregularly and poorly shaped, functional. Fruit a pear-shaped to nearly glo- and low branching. In Uganda TMrroeonthus hose capsule 2-3.5 cm long, 2-5-10bed, orange- o17iconus is an understorey tree with irregular, brown when ripe, opening near the apex, 2-5- low-branching hole, but in West and Central seeded. Seeds rounded to slightly triangular, Africa it may reach the upper canopy and de- 1.5-2.5 cm long, enclosed by a white or yellow velop a long, straight hole. In C6te d'Ivoire anI. Seedling with hypogeal germination; epl- flowering trees may be found throughout the coty1 4-6 cm long; first leaves alternate, SIm- year, but flowering is most abundant in March

, 572 TIMBERSl

to April, whereas fruits mature about 5 months transported by river. later. The fruits are eaten by animals, which Genetic resources Turroeonthus of neonus may disperse the seeds. However, seedlings are is widespread in West and Central Africa, but usually found in the close surroundings of it is uncommon in many regions within Its dis- mother trees tribution area. It is included in the IDCN Red Ecology Turroeonthus o17iconus occurs in list of threatened species as vulnerable because lowland evergreen forest and moist semi- it is exploited at moderate levels for its timber deciduous forest, often in humid localities and is becoming rarer in several regions where along streams and on the margins of poorly it was formerly common. However, it is still drained meadows and swamps. In Uganda it common locally in Ghana and C6te d'Ivoire. In grows up to 1500 in altitude. For Liberia it is Ghana it was rated as a 'pink star species', reported to prefer sandy soils exploited at a rate of less than 50% of the sus- Propagation and planting Natural regen- tainable cut, hence not at a rate to cause con- eration may be abundant near mother trees. cern. The 1000-seed weight is about I kg. The viabil- Prospects The wood of firroeonthus of neo- ity of seeds is very short and they must be itus is in high demand, especially for furniture sown immediately after collection. When fresh and decorative veneer. The often poor shape seeds are sown the germination rate may be up and small size of the hole is a serious draw- to 80% in 5-7 weeks. Young seedlings require back, as Is its very scattered occurrence in moist soiland quite deep shade. Seedlings need many regions More information Is needed on 10 months to reach 10 cm in height, and they growth rates and ecological requirements to have to stay in the nursery for at least one develop proper management strategies for sus- year, until the first compound leaves appear. tainable exploitation in natural forest. Wildlings are occasionally collected from the Major references ATIBT, 1986; Aubr6ville, forest for planting, but they are very suscepti- 1959a; Bolza & Keating, 1972; BurkiU, 1997; ble to drought. CIRAD Forestry Department, 2003; Farmer, Management 7'117roeonthus gifteditus usually 1972; Phongphaew, 2003; Siepel, Poorter & occurs scattered and in low densities in the Hawthorne, 2004; Vivien & Faure, 1985; Voor- forest. In south-western Cameroon the average hoeve, 1979 volume of holes larger than 60 cm in diameter Other references Adjanohoun at al. (Edi- is 0.12 mama. However, in West Africa it is tors), 1988; African Regional Workshop, 1998g; locally common, e. g. in Ghana, but also in C6te Akam at a1. , 2006; de Koning, 1983; de in d'TVoire, where in some coastal forests up to 5 Mensbruge, 1966; Hawthorne, 1995; Inside- trees with a bole diameter of more than 40 cm Wood, undated; Irvine, 1961; Katende, Birnie have been recorded per ha. The tree can be & Tengnas, 1995; Koeppen & Kukachka, 1961; managed by coppicing and pollarding Neuwinger, 2000; Normand & Paquis, 1976; Diseases and pests Roots of young plants in Odur0, 2003; Pauwels, 1993; Pennington & C6te d'Ivoire have been reported to be infected Styles, 1975; Saville & Fox, 1967; Staner & with the nematode Hylo, terno iuorense, causing GIIbert, 1958; Styles & White, 1991; Tatsimo at necrosis of tissue which may adversely affect a1. , 2005; Taylor, Cadet & Luc, 1978; Tayman the growth. at a1. , 2006 Harvesting For trees in natural forest the Sources of illustration Voorhoeve, 1979; minimum felling diameter at breast height Wilks & TSSemb6, 2000 (dbh)is 60 cm in C6te d'TVoire, 70 cm in Ghana Authors F. W. Owusu and 80 cm in Liberia. Logs may have brittle heart; some caution is needed during felling operations. VanlHADiFFUSA S. Dransf. Yield With a minimum felling limit of 70 cm dbh, the annual allowable cut of Turroeonthus Protologue Kew Bull. 53(2): 381 (1998). of neonus in Ghana was estimated at 23,000 ms Family Poaceae (Gramineae) in 1997 Origin and geographic distribution Voltho Handling after harvest After felling, logs diffuso is endemic to Madagascar. should be removed immediately from the forest Uses In Madagascar the stems are used for and rapidly converted or be treated with pre- light construction, for making fences, and for servatives, because the wood is very liable to carrying and storing water. Formerly they blue stain. Logs float in water and can be were used for making a traditional musical

, VAUGHANIA 573 instrument, the tube zither known in Mada- requirements, propagation, cultivation and gascar as 'vanha'. Nowadays this instrument is yield. They do not seem bright, however, be- made from other bamboo species with longer cause Voltho diffuso does not form clumps and internodes. The split and flattened sterns are does not respond wento excessive harvesting of woven into mats used in the construction of the stems walls and roofs Major references Bystriakova, Kapos & Properties The shoots contain cyanide Lysenk0, 2004; Dransfield, 1998 Botany Bamboo with elongated, necked rhi- Other references Clayton, Harman & Wil- 20mes; stem (CUIm) solitary, up to 10 in talland namson, 2002- a; Tan, 2006. up to 10 cm in diameter, woody, hollow, Triter- Authors M. Brink nodes 40-60 cm long with walls 2-3 mm thick; young shoots pale green, with dark brown hairs; leafy branches 20-30 cm long. Leaves VAUGHANIADIONAEIFOLIA S. Moore alternate, simple; sheath up to 20 cm long and up to 21 cm wide at base, dark brown hairy Protologue Journ. Bot. 58: 188 (1920) when young, glabrescent; 11gule consisting of Family Papilionaceae (Leguminosae - Papil- long bristles; blade lanceolate to ovate- ionoideae, Fabaceae) lanceolate, 5-8 cm x I-1.5 cm, glabrous or with Origin and geographic distribution Vou- scattered white or brown hairs. Inflorescence a ghonio dionoeifolio is endemic to north-western contracted raceme-like panicle, 3-6 cm long and northern Madagascar. Spikelets more or less cyllridrical when young, Uses The wood is used for house construction 12-35 mm long, pale green, conslsting of 5-6 and cattle enclosures glumes and I fertile noret; lower glume c. 9 Properties The heartwood is brown and min x 2 min, 15-veined, second glume 9-11 mm heavy. x 2-3 mm, 17-veined, other glumes 15-20 mm Botany Deciduous shrub or smalltree up to x 6-8 mm, 23-27-veined, lemma 18-26 mm x 10 in tall; hole up to 40 cm in diameter; bark 8-12 mm, 29-41-veined, palea 14-24 mm x 4-8 fissured, greyish brown; branches with thick mm, 2-keeled, grooved, with 8-11 veins at ei- and robust short shoots. Leaves alternate, clus- ther side of the groove; 10dicules 3, I-3 min tered at the ends of short shoots, I-foliolate; long; noret with 6 stamens c. 15 min long, an stipules fused, triangular, 2-5 mm long, persis- ovary with style c. 5 mm long and 3 stigmas tent and covering the short shoots; petiole flat- Fruit an oblong caryopsis (grain) c. I cm long, tened and winged, up to 4 cm long and 1.5 cm flattened when dry, smooth, wallpapery, easily wide; leaflet ovate to circular, 1.5-5 cm x 1.5-4 removed. cm, rounded or slightly notched at apex, with Voltho comprises 2 species, but the second spe- minute 2-branched hairs on both sides. Inno- cies, Voltho perrieri(ACamus) S. Dransf. , has rescence an axillary sessile raceme I-2 cm only been collected once, in northern Madagas- long, few-flowered. Flowers bisexual, papil- car. ionaceous; calyx shallowly cup-shaped, 2-3 min Ecology Voltho diffuso was formerly re- long, with small, narrowly triangular teeth; stricted to primary forest up to 700 in altitude, corolla asymmetrical, pale purple to violet or but it has become more widespread and now reddish, standard circular, 10-12 mm long, also occurs on open hills and in valleys, often wings distorted, keel twlsted and spiralIy together with Rouenolo mudogoscorie, 1818 curled; stamens 10, fused into a sheath for Sonn. about two-thirds of their length, twisted and Management After harvesting the stems are curved upwards; ovary superior, shortly hairy, split on one side and flattened, and they are I-celled, style curved upwards. Frult a linear- woven into large panels used for walls and cylindrical pod 2.5-4 cm long, sparsely hairy, roofs dehiscing with 2 spiralling valves, 6-10-seeded. Genetic resources and breeding Voltho Seeds ellipsoid, 3-4 mm long, pale brown diffuso seems not to be threatened by genetic Voughonio dionoeifo!to usually flowers when it erosion as it is becoming more widespread in is leafless, shortly before the new leaves de- Madagascar. velop. Prospects Voltho diffuso is a IOCaUy useful youghunio comprises 11 species and is con- wild source of construction material. To assess fined to Madagascar. It is related to the much the prospects for domestication of this bamboo, larger genus Indigofero, which differs in its more information is needed on its ecological straight wings, keel and stamens, longer sta-

. 574 TIMBERSl minalsheath and free stipules. SUIt in peptic ulcers. In South Africa roots are The wood of youghonio clotse!it (Drake) Du used to treat infertility in women and menor- Puy, Labat & Schrire, a shrub or smalltree up rhagia. The roots are also used in traditional to 10 in tall with 7-Ile15) leaflets per leaf, is medicine to treat cardiac pain, colic and innu- used for house construction in southern Mada- enza. Pulverized leaves are applied externalIy gascar. youghonio CIOiseltiis classified as unl- against headache. In Reunion and Mauritius nerable in the IUCN Red list of threatened leaf decoctions and infusions are used to wash species. The stem of Voughonio interrupto Du wounds and sores, against pulmonary infoc- Puy, Labat & Schrire, a shrub up to 3.5 in tall tions, rheumatic pains, fever, Influenza, stom- with generally 3-5 leaflets per leaf, is used for ach-ache and amenorrhoea, and as astringent poles in house construction in southern Mada- Root and stem decoctions serve as an anodyne gascar for women during child delivery. In Reunion Ecology Voughonio dionoeifo!to occurs in the tree is planted as an ornamental and to deciduous woodland on sandy soils, up to 200 restore degraded environments. in altitude. Properties The heartwood is white to pale Genetic resources and breeding Although greenish brown, and indistinctly demarcated Voughonio dionoeifolio is restricted to north- from the up to 12.5 cm wide sapwood. The western and northern Madagascar, it is fairly grain is usually straight, texture fine and even. widespread there, with several protected areas Growth rings are distinct. located within its range. Therefore, it does not The wood is fairly heavy to very heavy, with a seem to be immediately endangered by genetic density of (740,820-900(-1060) kg/ms at 12% erosion. However, the number of corrections in moisture content. It is hard, tough and elastic. herbaria is limited and therefore the species It should be air dried slowly to avoid serious maybe uncommon surface checking. The rates of shrinkage from Prospects It is unlikely that youghonio green to oven dry are 5.5% radial and 9.3% dionoeifolio and other Voughonio spp. will tangential. Once dry, the wood is moderately become more important in the future as timber stable in service trees because they are too uncommon or too At 12% moisture content, the modulus of rup- smallin size. ture is 134-139 N/min2, modulus of elasticity Major references du Puy at a1. , 2002. 16,300-17,000 N/min2, compression parallel to Other references du Puy & Labat, 19981; grain 64-67 N/min2, shear 18-19 N/min2 du Puy, Labat & Schrire, 1994; Schatz, 2001. Janka side hardness 11,030-11,200 N and Authors R. H. M. J. Lemmens Janka end hardness 12,320-12,450 N The wood is rather difficult to saw because of its hardness;it is recommended that it is sawn VEPRISLANCEOiATA (Lain. ) G. Don before drying. It works and planes readily, and can be finished to a smooth surface. The gluing Protologue Gen. hist. I: 806 (1831). and turning properties are satisfactory. The Family Rutaceae wood Is not durable. Chromosome number2n= 72 Leaves and branches contain alkaloids and Synonyms Vepris undu!ato (Thunb. )I. Verd 11monoids. Leaf and root bark extracts showed & GASm. (1951), nom. illegit moderate in-vitro antiplasmodial activity Vernacular names White ironwood (En). Aqueous leaf and stem extracts showed moder- Bois patte poule, patte poule sans piquant (Fr). ate antibacterial and antifungal activities Origin and geographic distribution Vepris Botany Evergreen shrub or sinaU to medium- loneeoloto occurs from Kenya along the coast sized tree up to 20(-24) in tall, but often much south to South Africa, and additionally in smaller; hole of larger trees straight and cylin- Reunion, Mauritius and Rodrigues drical, up to 150 cm in diameter; bark surface Uses The wood is used for beams, tool han- grey to purplish grey, usuaUy smooth; crown dles and implements. It is suitable for heavy rounded; twigs glabrous. Leaves alternate, 3- construction, flooring, shipbuilding, vehicle foliolate; stipules absent; petiole I-5(-7) cm bodies, furniture, cabinet work, mine props, long, slender; leaflets sessile, narrowly ellipti- sporting goods, toys, novelties, precision cal, (3-)5-12 cm x (I-)1.5-3.5 cm, curieate at equipment, carving, vats and turnery. base, acute or obtuse at apex, margin often In Tanzania leaf and root decoctions are used wavy, glabrous, with numerous glandular dots, to treat malaria; however, high doses may re- pinnately veined with numerous lateral veins

. VEPRiS 575

Inflorescence a terminal panicle up to 12 cm a1. , 1994; Gunb-Fakim at a1. , 2005; Lavergne & long. Flowers unisexual, regular, 4-merous; Vata, 1989; Neuwinger, 2000; Sarrailh at al. , pedice1 I-3(-7) mm long; sepals united at base, 2007; Steenkamp, 2003; van Vuuren, Banks & c. 0.5 mm long; petals free, obovate or eUiptical, Stohr, 1978 c. 2 mm x I min, greenish yenow; male flowers Authors R. H. M. J. Lemmens with (7-)8 stamens shorter than petals and rudimentary ovary; female flowers with supe- nor, globose ovary, 4-celled, with sessile, disk- VEPRIS NOBILIS (Denle) Mziray shaped stigma, stamens rudimentary. Fruit a slightly depressed globose drupe 4-8 mm in Protologue Acta Univ. Upsal. , Symb. Bot diameter, slightly 4-lobed, black when ripe, Upsa1. 30(I): 74 (1992) glandular dotted, usually 4-seeded. Seeds Family Rutaceae slightly trigona1, 2-3 min long, black Synonyms Tecleo nobilis Dellle (1843). Vepris loneeoloto grows moderately fast. In Vernacular names Small-fruited tealea (En). southern Africa trees flower in December- Origin and geographic distribution Vepris March and fruits ripen a few months after nobilis occurs from the Central African Repub- flowering. In South Africa it has been reported Iic, Sudan, ETitrea and Ethiopia south to Zim- that porcupines eat the bark, which may kill babwe, and additionally in Saudi Arabia and trees. The fruits are eaten by birds, which dis- Yemen perse the seeds Uses The wood is used for poles and posts in Vepris comprises about 80 species, most of house building, fences, tool handles and uten- them in mainland Africa, about 30 endemic to sils, e. g. walking sticks, clubs, spear shafts, Madagascar, and I in India bows and spoons. The wood is excellent for Ecology Vepris loneeo!oto occurs in coastal turning and inlay work, and Is also suitable for evergreen thickets, also on sand along the heavy construction, flooring, joinery, shipbuild- beach and on dunes. In South Africa it is also ing, vehicle bodies, furniture, cabinet work, found more lnland in dry evergreen forest, mine props, sporting goods, agricultural jin- where it reaches its largest dimensions. In for- PIements, toys, novelties and vats. It is used as ests in the southern parts of South Africa, firewood and for charcoal production. Vepris loneeo!oto is locally a doinlnant canopy The fruits are edible. The flowers are a source tree. In the Mascarene islands it occurs in low- of nectar for honeybees. The tree is occasionally land forest. planted for improving the soil by Its leaf litter, Management Vepris loneeoloto can be pro- and as a shade and amenity tree. The leaves pagated by seed. Seedlings transplant well. are used in a vapour bath to treat fever, and Trees can be managed by pruning. Logs should leaf and root decoctions are drunk to treat be removed from the forest soon after felling pneumonia, rheumatism and itching. The roots because they tend to split severely. serve as an arithelmintic and for treatment of Genetic resources and breeding Vepris pneumonia Pounded root bark is applied to lanceoloto does not seem to be immediately syphilitic ulcers, whereas the stem bark is threatened by genetic erosion because of its taken as an expectorant. Bark and leaves are wide habitat adaptability. However, in regions used as an analgesic. Roots and twigs are used where the trees reach larger sizes, which is the as toothbrushes. case in certain areas of South Africa, there has Properties The heartwood is creamy white, been overexploitation in the past. This may often with a dark brown core, and rather Iridis- mean that particularly superior timber geno- tinctly demarcated from the slightly paler sap- types are threatened. wood. The grain is usuaUy straight, texture Prospects Vepris loneeoloto provides a good- fine and even. Growth rings are distinct. Fresh quality timber, but more research is needed on wood often has an unpleasantsmell. tree development in relation to growing condi- The wood is fairly heavy, with a density of tions. It has prospects as an ornamental tree. 800-880 kg/in3 at 12% moisture content. It is Major references Behr, 2004; Bolza & hard and tough but fissile. It should be air Keating, 1972; Gunb-Fakim, Gu6ho & Bis- dried slowly to avoid serious deformation. At soondoya1, 1997; Palmer & Pitman, 1972-1974; 12% moisture content, the modulus of rupture Takahashi, 1978 is 134 N/min2. Other references Beentje, 1994; Coates The wood is rather difficult to saw and work, Palgrave, 1983; Gessler at a1. , 1995; Gessler at but it can be finished to a smooth surface. The 576 TIMBERSl

polishing and varnishing properties are satis- side tree. Vepris grushensis is classified as vul- factory, but the nailing properties are poor; nerable in the IUCN Red list. pre-boring is necessary in nailing. Steam bend- Vepris domellii(PICSerm. ) Kokwaro (synonym: ing gives good results. The wood is moderately Diphosio domellii PICSerm. ) is endemic to durable, being fairly resistant to termites, but southern Ethiopia, where it is a small under- liable to Lyctus and marine borer attacks storey tree up to 15 in tall in moist montane Tests In rats and mice ^showed anti- forest. It differs from Vepris nobilis by its OPPo- Inflammatory, analgesic and antipyretic activi- site leaves and 2-10bed fruit. The tough wood is ties of leaf extracts, without toxic effects, and used for furniture and agricultural imple- the triterpenoid Iupeol was isolated as the merits, and as firewood. The fruit is edible main active compound. Several quinoline alka- Vepris glunduloso (Hoyle & Leakey) Kokwaro, 101ds have also been isolated from the leaves, a shrub or small tree up to 7 in tall and en- as well as axane and OPPositane sesquiter- demic to Kenya, resembles yepris 810meroto, penes. but differs in its stalked and larger leaflets and Botany Evergreen shrub or smallto medium- fruits. Its wood is used for tool handles. yepris sized tree up to 20(-25) in tall; hole branchless glanduloso is classified as endangered in the for up to 7.5 in, often crooked, up to 60 cm in TUGN Red list. In 1995, fewer than 200 adults diameter; bark surface grey, smooth or finely were counted grooved; crown spreading; twigs glabrous. Vepris glomeroto (F. Hoffm. ) Engl. (synonym Leaves alternate, 3-foliolate; stipules absent; reeleo pi!o80 (Engl. ) I, verd. ) occurs from petiole 1.5-6(-8) cm long, terete; petiolules up southern Sudan and Ethiopia south to Tanza- to I cm long; leaflets elliptical to oblong- nia. It Is a shrub or small tree up to 7.5 in tall elliptical, 5-15(-18) cm x 1.5-4(-5.5) cm, Its wood is used for similar purposes as that of curieate at base, acute to acuminate at apex, Vepris nobilis, e. g. for house poles, sticks and margin entire, glabrous, with numerous glan- clubs. A root decoction is taken as a tonic and dular dots, pinnately veined with numerous to treat malaria, and a bark decoction to treat lateral veins. Inflorescence an axillary or ter- cardiac pain. The vapours of a root bark decoc- minal panicle up to 15(-21) cm long, glabrous tion are applied in cases of eye complaints, and Flowers unisexual, regular, 4-meTous, fragrant; ground roots are taken against hookworm. The pedicel up to 2 min long; sepals united, up to I foliage Is browsed by goats and camels. The min long, with small ovate lobes; petals free, fruit is edible narrowly elliptical, c. 4 min x 1.5 min, whitish Vepris grondifolio Engl. is a shrub or small yellow; male flowers with 4 stamens up to 5.5 tree up to 11 in tall occurring from Cameroon mm long and rudimentary ovary; female flow- east to Kenya and south to Angola and Zambia, ers with superior, globose ovary I-1.5 mm in and differs from yepris nobilis in its hairy in- diameter, I-celled, with short style and disk- norescences and smaller flowers. The hard and shaped stigma, stamens rudimentary. Fruit an tough wood is used for utensils, tool handles, obovoid drupe 6-8 mm x 5-6 mm, orange-red, bows, sticks, poles and pegs. It is also used as smooth, glabrous, I-seeded. Seed ovoid, 5-6 fuelwood. The tree is planted as an ornamental mmlong shade tree and roadside tree Vepris nobilis grows moderately slowly. In Vepris hanoitgensis (Kokwaro) Mziray (syno- southern Africa trees flower in August- nym: Teeleo honongensis Kokwaro) is a me- December, in Kenya in January-April and dium-sized tree up to 20 in tall, occurring in June-December, and fruits ripen I-2 months Kenya and Tanzania. Its wood is used for after flowering poles, tool handles, bows and spoons, and as Vepris comprises about 80 species, most of fuelwood. Vepris hurtongensis is planted as a them in mainland Africa, about 30 endemic to roadside tree Madagascar, and I in India. The wood of sev- Vepris inorogorensis (Kokwaro) Mziray (syno- eral other Vepris species is occasionally used in nym: Diphosio ino70gorensis Kokwaro) Is a East Africa: smalltree up to 10 in tall, endemic to Tanza- Vepris urushensis Kokwaro is a small to me- nia. Its wood is used for poles, tool handles, dium-sized tree up to 17 in tall, endemic to spoons and bows. It is also used as fuelwood northern Tanzania. Its wood is used for poles, Vepris inorogorensis is planted as an ornamen- tool handles, walking sticks, bows and spoons. talshade tree and roadside tree It is also used as fuelwood. Vepris orL, sheitsis is Vepris simplicifolio (Engl. ) Mziray non Endl. planted as an ornamental shade tree and road- (synonym: Teeleo simpltcifolio (Engl. ) TVerd. )

. VIGUIERANTHUS 577 is an illegitimate name used for a small I- Other references A1 Rehaily at a1. , 2003; A1 foliolate tree up to 10(-20) in tall, occurring In Rehaily at a1. , 2001; Bekele-Tesemma, Birnie & Ethiopia, Kenya and Tanzania. Its wood is very Tengnas, 1993; Hamill at a1. , 2003; Katende, similar to that of Vepris nobilis, and used for Birnie & Tengnas, 1995; Kokwar0, 1993; similar purposes In traditional medicine a Lovett at a1. , 2007; Neuwinger, 2000; Sommer- bark decoction is drunk to treat chest coin- Iatte & Sommerlatte, 1990; Wimbush, 1957 plaints and a root decoction to treat stomach- Authors R. H. M. J. Lemmens ache, backache, leprosy, gonorrhoea and brucellosis. Leaves and twigs are used for treating pleurisy. A leaf decoction is taken VIGuiERANTHUS KONY (R. Vig. ) Viniers against pneumonia, leaf ash is applied exter- nany against leprosy, and fruits are chewed to Protologue DUPuy, Legum. Madagascar: 277 relieve toothache. Twigs are used as (2002) toothbrushes. Family Mimosaceae (Leguminosae - Mimo- Vepris sto!zii TVerd. is a small tree up to 15 in soldeae) tall, distributed in Central and East Africa and Synonyms Colliondro kony R. Vig. (1949). in Angola. Its wood is used for poles in build- Origin and geographic distribution Vigttie- ing, spoons, handles, bows and as firewood 70nthus hony is endemic to the eastern part of Vepris storeii is planted as an ornamental central Madagascar shade tree and roadside tree Uses The wood is used for joinery Ecology Vepris nobilis occurs in evergreen Properties The wood is yellow-grey and forest, often together with Podocorpus and dense JuniperMs. and in riverme forest and wood- Botany Smallto medium-sized tree up to 18 land, at 900-2700 in altitude in East Africa, in tall; hole up to 30 cm in diameter; bark yel- but in southern Africa also at lower altitudes lowish grey; young branches grey, glabrous Management There may be abundant natu- Leaves alternate, bipinnately compound with a ral regeneration under parent trees. Pre- single pair of pinnae; stipules leathery, persis- treatment of seed is not necessary before sow- tent; petiole 4-12 min long, with winged mar- ing, but the germination rate of fresh seed is gins and gland at apex on upper side; pinna generally low and seeds lose their viahinty axis I-3.5 cm long, with winged margins; leaf- within a few months. One kg contains about lets 11-19 per pinna, alternate to opposite near 20,000 seeds. Wildlings are sometimes used for apex, sessile, oblong-rhombic to oblong- propagation. Trees can be managed by pruning obovate, up to 17 mm x 6 mm, asymmetrical at and pollarding base, rounded to acute at apex, glabrous. Inflo- Genetic resources and breeding VeprLs rescence an axillary raceme 2.5-6 cm long, nobilis is widespread, but within Its area of solitary or in clusters, many-flowered; peduncle distribution it is often uncommon and it has 2.5-3.5 cm long. Flowers bisexual, regular, 4- been subject to overexploitation in several re- 5-merous; calyx cup-shaped, c. 0.5 mm long, gions, either for its timber, e. g. throughout leathery; petals fused at base, (5.6-7.5 mm Uganda, or for medicinal purposes, e. g. in sev- long, pale greenish yellow; stamens numerous, eralregions in Kenya. Therefore, some caution fused at base, 14.5-18 min long; ovary supe- is needed to avoid genetic erosion nor, stalked, hairy, I-ceUed, style long and Prospects Vepris nobilis provides timber of slender. Fruit not known fair quality, but its production is hampered by Vigttieronthus comprises about 23 species, 18 the often poor shape and small size of the hole of them endemic to Madagascar, the remaining It is good for turnery and inlay work where species in tropical Asia. The wood of several smaller dimensions are acceptable. Tests in Viguierunthus species is used in Madagascar animals confirmed several of the claimed me- The wood of Viguieronthus cylindricostochys dicinal properties, such as analgesic, antipy- Viniers, a smalltree up to 15 in tall from cen- Tetic and anti-inflammatory activities. This tral and eastern Madagascar, is used for con- deserves further research on possibilities for struction and as firewood, that of Vtgttie- drug development rortthus longirocemosus Viniers, a medium- Major references Bolza & Keating, 1972; sized tree up to 20 in tall from eastern Mada- Maundu & Tengnds, 2005; Mbuya at a1. , 1994; gascar, is used in house building, and that of Takahashi, 1978; World Agroforestry Centre, Viguieronthus perui!lei (Drake) Viniers, a undated. smalltree up to 12 in tall from northern and 578 TIMBERSl eastern Madagascar, is also used for construc- vats, furniture, stools, carving, tool handles, tlon gunstocks, bowls, spoons, drums, guitars and Ecology Viguieronthus Aony occurs in humid beehives. It is also suitable for light construc- evergreen forest, from sea-level up to 1000 in tion, light flooring, joinery, interior trim, ship- altitude. It is locally common building, vehicle bodies, agricultural imple- Genetic resources and breeding The merits, toys, novelties, boxes, crates, matches, comparatively small distribution area of veneer, plywood, hardboard, particle board, Viguieronthus hony makes it liable to genetic wood-wool and pulpwood. The wood is used as erosion, although it is apparently locally coin- firewood and for charcoal production. mon. There are only a few protected forests in Cooked young leaves are eaten as a vegetable this area. or in sauces. The blackish pulp of the fruits is Prospects Viguieronthus hony and other Vi- edible and sweet, and eaten raw. It is often guteronthus species will remain of little local used to makeJam. Abeverageis madefrom the Importance for their timber, being either fruitjuice, and boiled fruits are the basis for an smaller-sized trees or uncommon alcoholic liquor and wine. The seeds inside the Major references du Puy at a1. , 2002. fruit stone are also edible Other references Lewis at a1. , 2005. Vitex doniono has numerous applications in Authors R. H. M. J. Lemmens traditional medicine. Leafsap is used as an eye drop to treat conjunctivitis and other eye coin- plaints. A leaf decoction is applied externalIy VITEXDONIANASweet as a galactagogue and against headache, stiffness, measles, rash, fever, chickenpox and Protologue Hort. brit. , ed. I, 2: 323 (1826). hemiplegia, and internally as a tonic, anodyne Family Verbenaceae (APG: Lamiaceae) and febrifuge, and to treat respiratory diseases Chromosome number2n=32 Pastes of pounded leaves and bark are applied Synonyms Vitex curieoto Thorni. (1827), Vitex to wounds and burns. Leafinfusions are added cienfoowskii Kotschy & Peyr. (1867), Vitex po- to alcoholic drinks to make them stronger. A chyphyllo Baker (1900) root decoction Is administered orally to treat Vernacular names Black plum, West Mri- ankylostomiasis, rachitis, gastro-intestinal can plum (En). Prunier noir, koro (Fr). Getona disorders and jaundice, and as an anodyne (Po). Mfudu, infuru, infuu (Sw) Powdered bark added to water is taken to treat Origin and geographic distribution Vitex colic, and a bark extract is used to treat stom- doniono is extremely widespread in tropical ach complaints and kidney troubles. The bark Africa, occurring from Senegal east to Somalia is also used against leprosy and liver diseases, and south to South Africa; also in Comoros and and to control bleeding after childbirth. Dried Seychelles. It is occasionally cultivated else- and fresh fruits are eaten against diarrhoea, where, e. g. in Mauritius and as a remedy againstlack of vitamin A and Uses The wood is popular for house building, B. The twigs are used as chewing sticks for teeth cleaning The blackish extract obtained by boiling leaves, bark, roots and/or fruits is used as ink and as a dye for clothes. The flowers serve as source of nectar for honeybees. Cattle browse the foliage Vitex doniono is planted as an ornamental shade tree. It contributes to the improvement of soilfertility by litter production Production and international trade Vitex o doniono timber appears to be traded mainly in Q local markets and in small amounts. Very smallvolumes have been traded from Gabon to Europe under the name 'evino'. Fruits are also ,, , traded on local markets and are locally coin- o mon in the fruiting season. The leaves are traded locally as a vegetable. Properties The heartwood is creamy white Vitex doniono - wild to pale brown, yellowish brown or greyish

. ViTEx 579 brown and indistinctly demarcated from the boons (Popio hornodryos grillbis). The presence 2.5-6 cm wide sapwood. The grain Is straight of progestogen-like compounds in the fruit has to wavy or interlocked, texture moderately fine been suggested as the probable cause. The seed to moderately coarse. The wood resembles oil has high iodine and low saponification val- teak. It is medium-weight, with a density of ues and can be used for skin cream, resin and 430-620 kg/ina at 12% moisture content, and paint production. Dried seeds yLeld about 30% soft. The wood air dries fairly easily, with little oil deformation although there is a tendency to Oral and intravenous administration of an cup. The rates of shrinkage from green to oven aqueous stem bark extract produced a dose- dry are 1.8-3.1% radial and 5.5-7.0% tangen- dependant hypotensive effect in both nor- tial, and from green to 12% inolsture content inotensive and hypertensive rats. The results about 1.1% radial and 3.3% tangential. Once of tests on rats suggest that an aqueous extract dry, the wood is stable in service of the stern bark is hepato. protective. The At 12% moisture content, the modulus of rup- aqueous extract of chewing sticks made of Vitex ture is 41-129 N/min2, modulus of elasticity doniono purchased in the market exhibited 5000-6500 N/min2, compression parallel to strong activity against a wide spectrum of bar- grain 27-50 N/mm2, shear 7-9 N/min2, cleav- tena including medically and dentalIy relevant age 59 N/mm radial and 74 N/mm tangential, bacteria, although the extracts of chewing and Janka side hardness 3020 N. sticks from Goreiniu hold Heckel and Anogeis- The wood is easy to saw and work with hand sus leiocorpu (DC. ) Gulll. & Perr. had broader and machine tools. It often planes to a silky or and generally stronger effects. This supports furry surface due to the presence of interlocked the traditional use of these chewing sticks with grain. The wood nails well with little splitting, reported anticaries effect. From tests with hot but it does not always hold nails well. Veneer water extracts of the bark on uterine muscle of good quality can be produced, but the logs strip preparations, it was concluded that the are often too irregular to be suitable for rotary use of the bark to control postpartum bleeding peeling. The wood is often too soft for turnery may be justified. The traditional use of Vitex It is usuaUy not durable, although good dura- doniono against diarrhoea was supported in hinty has also been reported, especially resis- tests with aqueous methanol extracts of the tance against termites. The wood is moderately stem bark on perfused isolated rabbitjejunum resistant to impregnation with preservatives. and on castor oil-induced diarrhoea in mice The wood dust may cause dermatitis in work- Stem bark extracts were able to inhibit the ers growth of clinical isolates of Salmonello typhi, The fruits are acidic, the fruitjuice has a pH of Shigello dysenterioe and Escherichia coli, sug- about 4.5. Per 100 g edible portion, the coinpo- gesting that they may be valuable in the sition of the fruit pulp is: water 59.5-73.5 g, treatment of dysentery and other gastroen- energy 435 kJ (104 kcal), protein 0.6-0.8 g, fat tentic Infections 0.1-1.3 g, carbohydrate 27.5 g, fibre 1.3 g, Ca The annual quantity of dry litter produced by 20-47 ing, P 47 ing, Fe 2.0-4.5 ing, ascorbic Vitex doniono in northern Cameroon Is about acid 6-18 ing (Leung, Busson & Jardin, 1968). 200 g/in2. Jam prepared from the fruit showed no signifi- Description Dedduous smallto medium-sized cant difference in flavour, colour and overall tree up to 25 in tall; hole branchless for up to acceptability from a commercial plum jam, and 11 in, up to 90(-160) cm in diameter, often the Vitex doniono jam was even preferred for slightly fluted at base; bark surface greyish consistency and spreadability. The fruits are a white to pale greyish brown, fissured and scaly, good source of potassium and iron. The syrup inner bark yellowish white, darkening to made from the fruit pulp can be used instead of brown; crown rounded; young branches shortly other syrups as a nutritive sweetener. Wine hairy, glabrescent. Leaves opposite, digitately obtained from controUed fermentation had compound with (3-)5(-7) leaflets; stipules ab- 10.5% alcoholcontent, and wine obtained from sent; petiole 5-20 cm long; petiolules up to 2.5 spontaneous fermentation 5.0%. Several my- cm long; leaflets obovate to elliptical, 4-25 cm cotoxigenic yeasts and fungiwere isolated from x 2.5-10.5 cm, notched to rounded or shortly deteriorating fruits; they may cause a health acuminate at apex, entlre, leathery, nearly hazard to livestock. Consumption of large glabrous. Inflorescence an axillary cyme up to amounts of fruits causes a transient reduction 10 cm long and 16 cm wide, orange-brown in reproductive functioning in female olive ba- hairy; peduncle 2-7.5 cm long; bracts up to 6 580 TIMBERSl

simple perforation plates; (21: intervessel pits opposite); 22: intervessel pits alternate; (25 intervessel pits small(4-7 F1m)); 26: intervessel pits medium (7-10 I'm); (30: vessel-ray pits , \ 4 with distinct borders; similar to intervesselpits in size and shape throughout the ray cell); 31 vessel-ray pits with much reduced borders to 1.1 apparently simple: pits rounded or angular; 32 fly ';{) vessel-ray pits with much reduced borders to apparently simple: pits horizontal(scalariform, gash-like) to vertical(palisade); 42: mean tan- 3 gential diameter of vessellumina 100-200 prn; 46: S 5 vessels per square minimetre; 47: 5-20 vessels per square minimetre. Tracheids and fibres: 61: fibres with simple to minutely bordered pits; 65: septate fibres present; 69: fibres thin- to thick-walled. Axial parenchyma: 78: axial parenchyma scanty paratrachea1; (79: IF:'.'.~,!:::?*g, axial parenchyma vasicentric); 92: four (3-4) cells per parenchyma strand; 93: eight (5-8) ,".""'~"";'*, *.. , In;;^ cells per parenchyma strand. Rays: 98: larger I!,, IF~ ~. *.,,, I rays commonly 4- to 10-senate; 106: body ray

.. 2 cells procumbent with one row of upright and/or square marginal cells; 1/5: 4-12 rays per mm. Mineral Inclusions: 152: crystals of other shapes (mostly small); (154: more than Vttex doniono - I, tree hubit, . 2, port o1/10wer- one crystal of about the same size per cell or ing 670nch, 3, port offrutting brunch, . 4, fruit. chamber). Redrawn grid adopted by Ishoh Syomsudin on. Thiam, P. Baas & P. Derienne) Growth and development The growth rate mm long. Flowers bisexual, zygomorphic, 5- of Vitex dontono is moderate. In plantations in merous; pedicel up to 2 min long; calyx conical, northern C6te d'Ivoire, seedlings were on aver- 3-5 mm long, with short teeth, enlarging in age 70-90 cm tall after 3 years, the tallest ones fruit; corolla white to pale purple, tube 6-8 min reaching 170 cm. On good soils in southern long, curved, limb 4-10bed, lobes c. 3 mm long Burkina Faso early growth is a bit faster. In and middle lower lobe up to 4.5 min long; sta- drier regions of West Africa trees flower in the mens 4.1nserted in the corolla tube, 2 long and second half of the dry season or at the begin- 2 short; ovary superior, obovoid, 4-celled, style rimg of the rainy season. In Gabon they lose c. 7 mm long. Fruit an obovoid to oblong- their leaves in December-February and flower ellipsoid drupe 2-3 cm long, purplish black, in March-May. The flowers are commonly vis- fleshy, with woody, 4-celled stone, up to 4- ited by bees and sunbirds. In Gabon fruits start seeded. Seeds without endosperm to ripen in August. They are eaten by monkeys, Other botanical information Vitex coin- gorillas, chimpanzees and elephants, which prises about 150 species and is pantropical probably disperse the seeds. In Tanzania fruits with a few species in temperate regions. Ap- ripen in April-July, in Zambia in April- proximateIy 60 species occur in tropical Africa, September. of which Vitex doniono is most widespread. Ecology Vitex doniono occurs in a variety of The vanability of Vitex doniono is remarkable, habitats, from forest to savanna, often in wet regarding its morphology as well as its habitat localities and along rivers, and on termite choice, and biosystematic research is war- mounds, up to 2000 in altitude. It occurs in ranted. regions with a mean annual rainfall of 750- Anatomy Wood-anatomical description OAWA 2000 mm. It is most commonly found on allu- hardwood codes) vial soils. In Central Africa it is often the first Growth rings: (I: growth ring boundaries dis- species to establish when gallery forests evolve tinct); (2: growth ring boundaries indistinct or in low-lying areas in the savanna. absent). Vessels: 5: wood diffuse-porous; 13: Propagation and planting Vitex doniono

. VITEX 581 is propagated by seed or root suckers. The 2000; Bryce, 1967; CAB International, 2005; weight of 1000 fruit stones is about I kg Egbekun, Akowe & Ede, 1996; Higham at al. , Stones should be sown fresh after removal of 2007; InsideWood, undated; Kilani, 2006; Kris- the pulp and soaking in cold or warm water for tensen & Lykke, 2003; Ladeji & Okoye, 1993; 24 hours' In C6te d'TVoire, stones dipped into Ladeji & Okoye, 1996; Laden, Okoye & Uddoh, sulphuric acid 95% for 60 minutes and subse- 1996; Laden, Uddoh & Okoye, 2005; Leung, quently in water for 72 hours germinated after Busson & Jardin, 1968; Louppe & Ouattara, 26 days, but the germination rate was only 1993; Mapongmetsem at a1. , 2005; Neuwinger, 34%. Untreated fruits may take very long to 2000; Okigb0, 2003; Sambo & Maghembe, germinate; fire may accelerate germination 1995; Taiwo, Xu & Lee, 1999; Thiel at a1. , 1993. Stories may contain several seeds and several Sources of illustration Ruffo, Birnie & seedlings may germinate from one stone. Seeds Tengn6. s, 2002; Troupin, 1982. can be stored for up to I year at 3-5'C. Propa- Authors K. J. M. Ky gation by cuttings has been successful in Ma- lawi. Wildlings are sometimes also used for planting. Survival rates in plantations are VITE>CFISCHERIGiirke normaUy good, about 80-90% after 3 years Management Vttex doniono trees are often Protologue Bot. Jahrb. Syst. 18: 171 (1893) retained when clearing forest or savanna bush- Family Verbenaceae (APG: Lamiaceae) land. The tree can be managed by coppicing Chromosome number 271= 96 and lopping Synonyms Vitex keriiensis Turn11(19/5). Harvesting Fruits are usually collected from Vernacular names Meru oak, Kenya oak the wild, but in Nigeria they are sometimes (En). Mfuu, mrudu (Sw). also collected from trees planted on farms. Origin and geographic distribution Vitex Handling after harvest Felled logs are 11a- Itscheri occurs wild from DR Congo and south- ble to blue stain and should not be left un- ern Sudan east to Kenya and south to Angola treated in the forest for longer periods. Fresh and Zambia. It has been planted in the high- logs float in water and can be transported by lands of Kenya, Uganda and Tanzania, and in river. In Nigeria the fruit juice is fermented Cameroon using a traditional technology to produce wine. Uses The wood is used for furniture, coffin Genetic resources Vitex doniono is very boards, panelling and veneer. In Kenya it is in widespread and common in many regions. It is high demand for furniture, and is also popular not threatened by genetic erosion, although for making tool handles and oxen yokes. The locally populations may be under pressure. wood is suitable for light construction, flooring, Prospects Vitex doniono, a tree highly val- joinery, interior trim, shipbuilding, vehicle ued by people in various parts of Africa and for bodies, toys, novelties, boxes, crates, carving, various purposes, deserves protection Ln the turnery, draining boards, hardboard, particle wild and its domestication should be promoted board and pulpwood. It is used as firewood and In a study in Burkina Faso, Vitex doniono was amongst the 8 species most highly valued by the local population, especially as a source of construction timber, edible fruits and leaves to

make a vegetable sauce. It has Interesting . prospects for the development of modern phy- totherapeutical medicines, especially for treating gastro-intestinal diseases and for dental hygiene, traditional uses which have been confirmed by pharmacological research and which ,^ are mainly based on antibacterial actrvities. L. Major references Bolza & Keating, 1972;

Burki11, 2000; Latham, 2004; Maundu & .. Tengnas, 2005; Mbuya at a1. , 1994; Ruffo, Birnie & Tengn6. s, 2002; Takahashi, 1978; Q Verdcourt, 1992; von Maydell, 1986; World Agroforestry Centre, undated. Other references Agunu at a1. , 2005; mobo, fite" lischeri- wild 582 TIMBERSl

for charcoal production. ,~ -. The blackish pulp of the fruits is edible and 4 eaten raw. It has been recorded that the honey produced by bees visiting Vitex lischeriflowers , is of superior quality and fetches high prices in , I I the market. Vitex lischeriis planted as a shade ' I'\' tree for crops such as coffee and yam, and may be retained in maize and cassava fields. It is grown as an ornamental and windbreak. It ,,,,., , ';-,. I. -:;,"*,#. produces a useful mulch and serves as a soil Improver .;\"".,,;:-- 11\~..^I ,,,,, 1,1;;*4 ~,"' ' Production and international trade Meru ^-I I. - --.', oak timber appears to be traded only in local markets and in small amounts. In Kenya, .,*, \ I \\ ,/" where the wood is prized and in high demand, ,44 *..-.. t*$,: ~}' " " " ' the annual volume traded has been estimated ~, at about 400 ina. Fruits are traded on local markets on a smallscale . * * Properties The heartwood is pale grey-brown , and indistinctly demarcated from the sapwood, .!,' , with heartwood of older trees often dark ~~-, 2 stained and decorative. The grain is straight or wavy, texture coarse. The wood resembles teak, and often yellows after some time in service Logs are often rotten in the core. The wood is medium-weight, with a density of 430-570 kg/ina at 12% moisture content. It air dries Vitex lischeri - I, tree habit, 2, port o1/10uiering easily, with little deformation; the rates of brunch, . 3, flower, 4, inIructescence shrinkage are moderate. The wood is easy to Redruwn und udopted by Ishoh Syomsudin saw and work with hand and machine tools. It planes to a smooth surface, but a filler is minute teeth, enlarging in fruit; corolla white, needed for polishing and varnishing. It nails often tinged blue, tube 3-6 mm long, limb 2- well. The gluing properties are good, and ve- lipped, 3-5 min long, upper lip 2-fid, lower 3- neer of excellent quality can be produced. As- fid; stamens 4, inserted in the corolla tube, 2 sessments of the durability of the wood range long and 2 short; ovary superior, globose, 4- from nori-durable to durable. The heartwood is celled, style c. 6 mm long. Fruit an obovoid to moderately resistant to Impregnation with oblong-globose drupe I-2.5 cm long, blackish preservatives, the sapwood is permeable with whitish or greenish spots, fleshy, with Description Deciduous small to fairly large woody, 4-celled stone, up to 4-seeded. Seeds tree up to 35 in tall; bole branchless for up to without endosperm. 18 in, up to 180(-230) cm in diameter; bark Other botanical information Vitex coin- very thin, bark surface grey to dark brown, prises about 150 species and is pantropical smooth to rough and slightly fissured, inner wlth a few species in temperate regions. Ap- bark creamy yellow to pale brown; crown proximateIy 60 species occur in tropical Africa. rounded; young branches densely velvety Vite" henie, ,sis Turnllis often considered to hairy. Leaves opposite, digitately compound represent a separate species, distinguished with 5 leaflets; stipules absent; petiole 6.5-17 from Vite" lischeri by its larger tree size, more cm long; petiolules up to 4 cm long; leaflets shaggy indumentum, more lax inflorescence, obovate to elliptical, 5-19 cm x 3-10 cm, more distinctly toothed calyx and larger fruit rounded to acuminate at apex, entire, leathery, It occurs mainly in montane rainforest, densely hairy and glandular below. Innores- whereas Vitex lischeriis found in wooded cence an axillary cyme up to 12 cm long and 24 grassland and thickets. However, a biosys- cm wide, densely hairy; peduncle 6-14 cm long; ternatic study of Vitex heniensis and Vitex bracts up to 2 cm long. Flowers bisexual, zy- lischeri populations revealed no reason for gomorphic, 5-merous; pedice1 I-4(-12) min taxonomic separation. It was also demon- long; calyx campanulate, 2.5-4 min long, with strated that different populations are not re-

. ViTEx 583 productiveIy isolated such as bees. However, fruits can be produced Vite, c congo!errsis De Wild. & T. Durand is a by both self- and cross-pollination. In Kenya small to medium-sized tree up to 30 in tall, ripe fruits can be found from June to Novem- known from the Central African Republic and her, and fruit maturity coincides with leaf fall DR Congo. Its wood is suitable for similar pur- In Tanzania ripe fruits are available from April poses as that of Vitex lischeri; its density is to August. The seeds are dispersed by animals slightly higher (about 600 kg/in3 at 12% ino1s- such as monkeys and bombills that eat the ture content) and it is somewhat more durable fruits Anatomy Wood-anatointcal description (IAWA Ecology Vitex lischeri occurs in evergreen hardwood codes): forest, open woodland, wooded grassland and Growth rings: I: growth ring boundaries dis- thickets, up to 2100 in altitude. In Kenya the tinct. Vessels: (4: wood semi-ring-porous); (5: mean annual temperature in the distribution wood diffuse-porous); 13: simple perforation area of Vitex lischeriis 15-23'C, and mean plates; 22: intervessel pits alternate; 26: in- annual rainfall 1200-2000 min. On the slopes tervessel pits medium (7-10 pin); (30: vessel- of Mount Kenya it prefers deep sandy loam ray pits with distinct borders; similar to in- soils, but it is elsewhere locally common in tervessel pits in size and shape throughout the thickets on granite rocks and in wooded grass- ray cell); 31: vessel-ray pits with much reduced land on termite mounds. borders to apparently simple: pits rounded or Propagation and planting In montane angular; 32: vessel-ray pits with much reduced rainforest in central Kenya, natural regenera- borders to apparently simple: pits horizontal tion of Vitex lischeri occurs particularly in for- (scalariform, gash-like) to vertical mallsade); est gaps, and is poor in closed forest. Seeds for (33: vessel-ray pits of two distinct sizes or types propagation can best be collected when most in the same ray cell); 43: mean tangential di- fruits are still green while about 20% have ameter of vessellumina a 200 prn; 46: S 5 ves- turned brown. All fruits collected should have sels per square minimetre. Tracheids and fi- grown to the final size. After collection, fruits bres: 61: fibres with simple to inlnutely bor- are packed in gunny-bags. The pulp should be dered pits; 65: septate fibres present; 69: fibres removed within a few days, after which the thin- to thick-walled. Axial parenchyma: 78 stones are dried in the shade up to a minimum axial parenchyma scanty paratrachea1; (79: moisture content of 8.5%, and can be stored axial parenchyma vasicentric); (80: axial pa- subsequently. The germination rate of fresh renchyma anform); (81: axial parenchyma 102- seeds is 40-50% after 9 weeks. Fruit stones are enge-allform); 91: two cells per parenchyma usually used for propagation; one kg contains strand; 92: four (3-4) cells per parenchyma 2500-3000 stones. Several seedlings may ger- strand. Rays: (97: ray width I-3 cells); (98 inmate from one stone because it may contain larger rays commonly 4- to 10'seriate); 106 I-4 seeds. Soaking in cold water for 24 hours body ray cells procumbent with one row of up- improves germination. When the fruit pulp has right and/or square marginal cells; (110: sheath been removed and the stones dried in the cells present); 1/4: S 4 rays per min; 1/5: 4-12 shade, they can be stored for considerable time rays per mm (at least one year)ifthey are kept dry and cool. (M. Thiam, P. Baas & P. Derienne) Although seedlings are light demanders, they Growth and development Seedlings have tolerate shade. In the nursery they stillshowed simple leaves in the early stages and start to good growth at an Irradiance level of 19% of produce compound leaves after about 3 full sun, but higher levels of light enhanced months. Vitex lischeri trees grow moderately growth. Wildlings are sometimes also collected fast. In a plantation in Kenya, trees were on for planting. average 35 in tall 56 years after plantlrig. Management The firsttimber plantation was Maximum mean annual volume increment was established in the early 1950s on the slopes of reached 25-30 years after planting, and a Mount Kenya. In 1973 about 620 ha of planta- marked decline in volume increment was re- tions were recorded. The rotation for trees corded when trees were more than 35 years planted on farms for timber production is re- old. Trees of 50 years old may reach a hole di- corded to be 35 years, but in commercial timber ameter of 60 cm plantations with a stand density of 200 trees In Kenya trees flower from December to per ha, a cycle of 45-60 years seems more ap- March, before the long rainy season starts in propriate. Pruning is recommended to obtain a April. The flowers are pollinated by Trisects straight and clear hole. Trees tolerate coppic- 584 TIMBERSl

ing. They can also be lopped and pollarded, Other references Arap Sang & Munga, thus providing farmers with fuelwood while 1981; Beentje, 1994; Dale & Greenway, 1961; allowing the hole to reach marketable size. Dharani, 2002; Fernandes, 2005; InsideWood, Diseases and pests Seedlings in nurseries undated; Konuche, 1995; Pauwels, 1993; Wim- are susceptible to damping-offcaused by fungal bush, 1957. attacks. Parasitic plants of the genus Cuscuto Sources of illustration Dale & Greenway, can cause serious losses in nursery seedlings 1961 The fungus Armil!orig meIleo often attacks the Authors H. J. C. Thijssen tree in plantations resulting in a black resin on the stems and in decay accelerated by Trtcho- dermo sp. However, this disease is usually not VITEXGRANDIFOLIA Gtirke a serious threat and can be controlled by up- rooting affected trees at first thinning Protologue Bot. Jahrb. Syst. 18: 169 (1894) Genetic resources In general Vitex lischeri Family Verbenaceae (APG: Lamiaceae) is widespread and not uncommon in many re- Chromosome number2n= 32 gions in its distribution area. However, the Origin and geographic distribution Vitex populations in montane rainforest in central grondifolio occurs from Sierra Leone east to Kenya, where the holes are often straight and Gamero0n and Gabon reach large dimensions and are thus of high Uses The wood is used on a local scale for value in the timber trade, have been overex- light construction, drums and canoe seats. It is PIOited. These populations, referred to as Vitex suitable for light flooring, joinery, interior trim, heriiensis, are classified as vulnerable in the furniture, cabinet work, toys, novelties, ship- 2006 TUGN Red List. In several other habitats building, vehicle bodies, agricultural imple- and regions, Vite, c lischeriis also threatened merits, boxes, crates, veneer, plywood and because offorest clearing for human settlement pulpwood. The fruits are edible and used to and agriculture, e. g. near Lake Victoria and at make an alcoholic drink. In traditional medi- the foot of Mount Elgon cine the bark is used as a stomachic and to KBFRl and the Plant Conservation Pro- treat diarrhoea, bronchial complaints, rickets, gramme in Kenya maintain a living collection sores and fever. Leaves are used in medications of Vttex lischeri, as well as seed stocks. Re- against colic, infections of the umbilical cord, search showed that the bulk of genetic vana- toothache, rheumatism and orchitis. A tea tion in natural populations in Kenya is within made from the fruits is drunk as a tonic. The populations (96.5%), and therefore sampling black juice exuding from leaves heated over a from any of these populations would capture a fire has been used as ink sufficiently large genepool of the species for Production and international trade The breeding and conservation timber of Vitex grondifolio is used locally, al- Prospects Vitexftscheriis already planted in though there are also reports on export to agroforestry systems, but it has substantial Europefrom Garnero0n and Gabon potential for larger-scale planting, not only for Properties The heartwood is whitish to pale timber production, but also as a source of fire- brown, darkening on exposure to brown, and wood, edible fruits and honey, and as a useful indistinctly demarcated from the sapwood. The auxiliary tree. Efforts to promote planting and grain is usually straight, texture medium. The conservation of Vitex lischeri are hampered by wood has a density of about 490 kg/ina at 12% lack of information on its biological and eco- moisture content. The wood air-dries fairly logical characteristics. More research is needed easily, but has a tendency to cup; it may be on the variation in wood properties between reconditioned by steaming. The rates of different populations, and on the role of the shrinkage are moderately high: from green to ecological conditions on wood properties, the oven dry 4.0% radial and 6.4% tangential. Once results of which might be used in breeding pro- dried, the wood is moderately stable in service grammes. At 12% moisture content, the modulus of rup- Major references Menda, 1999; Bolza & ture is about 88 N/mm2, modulus of elasticity Keating, 1972; CAB International, 2005; Joker 7700 N/mm2, compression parallel to grain 35 & Mngulwi, 2000; Kigom0, 1985b; Maundu & N/min2, shear 8 N/mm2, cleavage 15 N/min and Tengias, 2005; Mbuya at a1. , 1994; Rutfo, Chalais-Meudon side hardness 2.5 Birnie & Tengnas, 2002; Verdcourt, 1992; The wood is easy to saw and work with hand WorldAgroforestry Centre, undated and machine tools. It generally planes to a

. ViTEx 585

smooth surface and finishes well. The wood o0n) the average timber volume was recorded nails well without splitting. Veneer of good at 0.65 matha. Logs are liable to blue stain at- quality can be produced. The wood is moder- tack and should be treated with preservatives ately durable and reportedIy resistant to ter- or removed from the forest soon after felling. mite attack. The sapwood is susceptible to Genetic resources and breeding There Lyetus beetle attack. The heartwood is moder- are no reasons to consider Vitex grondifo!to as ately resistant to impregnation by preserva- threatened by genetic erosion because it is tives; the sapwood Is fairly permeable. fairly widespread, at least locally common and Botany Evergreen shrub or smallto medium- also present in disturbed forest sized tree up to 20 in tall; hole branchless for Prospects Too little is known on ecology, up to 15 in but usually much shorter, up to growth rates and natural regeneration of Vitex 60(-120) cm in diameter, often sinuous and grondifolio to judge its prospects as a timber slightly fluted at base; bark surface grey to tree in sustainably managed forest. However, reddish brown or greenish yellow, finely scaly, the often poor shape and small size of the hole inner bark yellowish, rapidly darkening on seem to limit its possibilities for commercial exposure; young branches obtusely quadrangu- exploitation Iar, short-hairy or glabrous. Leaves opposite, Major references Bolza & Keating, 1972; digitately compound with 5(-7) leaflets; stip- Burki11, 2000; Sallenave, 1964; Takahashi, ules absent; petiole 9-20 cm long, stout; petio- 1978; Vivien & Faure, 1985. Iules up to 5(-10) mm long; leaflets obovate, Other references Adjanohoun & Ak6 Assi, 13-40 cm x 6-20 cm, acuminate at apex, en- 1979; Ako6gninou, van der Burg & van der tire, thin-leathery, glabrous. Inflorescence an Maesen, 2006; Aubr6ville, 1959d; de Koning, axillary, compact cyme up to 7 cm long, many- 1983; Hawthorne, 1995; Hawthorne & Jong- flowered; peduncle up to 5 cm long. Flowers kind, 2006; Huber, Hepper & MeIkle, 1963; bisexual, zygomorphic, 5-meTous; calyx 3-6 Raponda-Walker & Sillans, 1961 mm long, enlarging in fruit, with obscure teeth, Authors R. H. M. J. Lemmens finely hairy; corolla pale purple with yellowish limb, 15-20 min long, finely hairy; stamens 4, inserted in the corolla tube, 2 long and 2 short; VITEXMICRANTHA Gtirke ovary superior, globose, 4-celled, glabrous but hairy at apex, style slender, curved. Fruit an Protologue Bot. Jahrb. Syst. 18: 170 (1894) ellipsoid to globose drupe 1.5-2 cm long, yel- Family Verbenaceae (APG: Lamiaceae) lowish when ripe but later becoming black, Chromosome number2n=32 fleshy, with woody, 4-celled stone, up to 4- Origin and geographic distribution Vitex seeded. Seeds without endosperm. Seedling microntho occursfrom Guinea east to Ghana. with epigeal germination; hypocoty1 3-4 cm Uses The wood is used on a local scale for long, epicoty1 10-14 mm long; cotyledons thinly light construction, drums and writing boards. leathery, short-stalked; first pairs of leaves It is suitable for light flooring, joinery, interior simple trim, shipbuilding, furniture, cabinet work, In C6te d'Ivoire Vitex grondifolio trees can be toys, novelties, boxes, crates, veneer, plywood, found flowering almost throughout the year hardboard, particle board and pulpwood. The and fruits mature in March to May and Octo- fruits are edible and used to make an alcoholic her drink. In traditional medicine the leaves are Vitex comprises about 150 species and is pan- applied externalIy to treat onchocerciasis tropical with a few species in temperate re- (craw-craw), and various parts of the plant are gions. Approximately 60 species can be found probably used in similar ways to those of Vitex in tropical Africa. doniono Sweet. Ecology Vitex grondifo!to is often an under- Properties The heartwood is creamy white storey tree of lowland evergreen forest. In to pale yellowish brown, darkening on expo- many regions it is rather uncommon, e. g. in sure, and indistinctly demarcated from the Ghana and Gabon, but in other regions it may sapwood. The grain is straight to wavy, texture be locally common, especially in secondary for- moderately fine and even. The wood is fairly est, e. g. in C6te d'Ivoire and Cameroon lustrous. It has a density of about 520 kg/in3 at Management Seeds start to germinate 2-3 12% moisture content. The wood air dries fairly months after sowing. The germination rate easily. The rates of shrinkage are moderate: may be up to 90%. In forest near Ed6a (Camer- from green to oven dry 2.7-2.8% radial and

. 586 TIMBERSl

5.8-6.2% tangential. Once dry, the wood is orycuspis is used for house construction in Li- moderately stable to stable in service heria. It is similar to the wood of fitex micron- The wood is soft. At 12% moisture content, the tho, but the density is slightly higher (about modulus of rupture is 94-124 N/min2, modulus 580 kg/ina at 12% moisture content). The wood of elasticity 7000 N/mm2, compression parallel of Vitex phoeotricho Mildbr. ex WPIep. (syno- to grain 46-48 N/mm2, cleavage 20 N/mm and nym: Vitex rufo A. Chev. ex Hutch. & Dalziel), Chalais-Meudon side hardness 1.6 which occurs from Sierra Leone to C6te d'TVoire The wood is easy to saw and work with hand and from Gamero0n to Gabon, is also similar to and machine tools. It planes to a smooth sur. that of Vitex microntho. It is used for local face and finishes well. The wood glues satisfac- house construction, and additionally for tools, torny. Veneer of good quality can be produced, bowls and drums. The leaves and bark of Vitex but the shape of the hole is often too poor for phoeotricho, which is characterized by its rotary cutting. The wood is not durable and densely rusty brown hairy, obtuse or shortly liable to termite, powder post beetle and ina- acuminate and entire leaflets, are applied as rine borer attacks. The heartwood is moder. an antiseptic to ulcers. The fruits of Vitex Dry- ately resistant to impregnation by preserva- cuspis and Vitex phoeotricho are edible but tives; the sapwood is permeable. apparently not popular. Botany Evergreen, smallto medium-sized tree Ecology Vitex micronthu is often an under- up to 27 in tall; hole up to 60 cm in diameter, storey tree of lowland evergreen forest. It is often sinuous and slightly fluted at base; bark locally common. In Ghana it is strongly associ- surface grey to brown, fissured and scaly, inner ated with base-poor soils. bark yellowish brown, rapidly darkening on Management The 1000-seed weightis about exposure; young branches quadrangular, 500 g. Germination may start up to 5 months shortly hairy, glabrescent. Leaves opposite, after sowing. The germination rate Is often low. digitately compound with 3-5 leaflets; stipules In Sierra Leone Vitex microntho has been con- absent; petiole 3-6 cm long, slender; petiolules sidered a weed in production forest, and trees slender; leaflets obovate to oblanceolate, 9-12 have been killed with arsenite. cm x 1.5-3 cm, shortly acuminate at apex, Genetic resources and breeding Vitex toothed in upper part, papery, glabrous but microntho is fairly widespread and locally glandular dotted. Inflorescence an axillary, lax common and consequently not threatened by cyme, few-flowered; peduncle up to 20 cm long, genetic erosion. However, the ongoing reduc- slender. Flowers bisexual, zygomorphic, 5- tion and cutting up of the evergreen forest area merous; calyx c. 2 min long, with short teeth, in West Africa may endanger the species in the enlarging in fruit, finely hairy; corolla white future with pale purple limb, 6-7 min long, finely Prospects Too little is known on ecology, hairy; stamens 4, inserted in the corolla tube, 2 growth rates and natural regeneration of Vitex long and 2 short; ovary superior, 4-celled, gla- mrcrontho to judge its prospects as a timber brous. Fruit an ellipsoid drupe c. 1.5 cm long, tree in sustainably managed forest. However, purplish black, fleshy, with woody, 4-celled the often poor shape and small size of the bole stone, up to 4-seeded. Seeds without en- seem to limit its possibilities for commercial dosperm. Seedling with epigeal germination; exploitation. hypocoty1 3-4 cm long, epicoty1 4-8 min long; Major references Bolza & Keating, 1972; cotyledons leafy, shortly stalked; first pairs of Burki11, 2000; Kryn & Fobes, 1959; Saville & leaves simple Fox, 1967; Takahashi, 1978 In Cote d'Ivoire Vitex microntho trees flower Other references Aubr6ville, 1959d; de from February to April and fruits mature in Koning, 1983; de in Mensbruge, 1966; Haw- July. thorne, 1995; Hawthorne & Jongkind, 2006; Vitex comprises about 150 species and is pan- Huber, Hepper & Meikle, 1963; Sallenave, tropical with few species in temperate regions. 1955. In tropical Africa approximately 60 species can Authors R. H. M. J. Lemmens be found Vitex orycuspis Baker occurs from Sierra Leone to southern Nigeria, and possibly also in Cam- eroon, DR Congo and Angola. It resembles Vite" microntho, but generally has larger leaves and glabrous flowers. The wood of Vitex

. WIDDRINGTONIA 587

WIDDRINGTONIAWHYTEIRendle

Protologue Trans. Linn. SOC. London, Bot. 4 60 (1894). I^^ @ @ Family Cupressaceae Chromosome number2n=22 4 Synonyms Widdringtonio nodiftoro (L. ) POW- \ I tie va". whytei(Rendle) SIIba (1990) Vernacular names Mlanje cedar, inulanJe cedar, inIanje cypress (En) Origin and geographic distribution Wid- driftgtonio whyteiis endemic to Mount Mulanje in Malawi. It has been grown in plantations on

Zomba Mountain in Malawi and occasionally 3 .*. "" elsewhere -- f e , . * Uses Widdrtngtonio whytei, the national tree '* , of Malawi, has been one of the most important *,*. sources of softwood in Malawi for almost 100 I years. The wood has been used extensively for \ construction, furniture, panelling and fence .. posts. It has now become a scarce 'prestige I wood', highly valued for construction and boat -...- building. It is used for making hard-wearing roof shingles that weather to an attractive SII- . ver-grey, and it has been used for pencil manu- -...- 2 facture. It is used for making carvings, boxes , and furniture sold to tourists Production and international trade The Widdringtonio whytei- I, tree hubit, 2, ledding trade in the wood of Widdringtonio whytei has branchlet, . 3, brunch with seed cones, . 4, seeds. much diminished and is confined now to a local Redrown Grid udopted by Ishoh Syomsudin scale Properties The heartwood is yellow or pale not take paint well. Planed surfaces have a brown and is clearly demarcated from the nar- satiny sheen. The heartwood is durable and row pale sapwood. The grain is straight, tex- highly resistant to termites, borers and fungi ture fine and even. The wood is resinous and The sapwood is not susceptible to attack by has a persistent cedar-like odour. The density Lyetus beetles. of the wood is 530-610 kg/in" at 17.6% inois- The wood fibres are (3.8-)4.4(-5.1) mm long, ture content. The wood works easily. It does and the wood contains about 36% 11gnin, which is relatively high compared to other softwoods The wood can be used for papermaking, butthe high Iignin content could give problems during processing Steam-distillation of the sawdust gives a yield of about 10 inI essential oil per . 100 g moisture free material. The main constituents of the essential oil are thujopsene (32%), cedr01(14%), thujopsadiene (7%), wid- dr01(5%) and cuparene (4%) ,A Description Evergreen, monoecious, large tree up to 50 in tall; hole usually straight, L. branchless for up to 20 in, up to 150(-200) cm in diameter; outer bark grey-brown, smooth in .. young trees, thick, spongy, fissured and exfo}i-

o ating in long strips in older ones, inner bark red-brown; crown pyramldal, eventually Irregu- Iar or nat-topped; branches spreading or as- Waddrtngtonio whytei- wild cending. Leaves decussately opposite on small- 588 TIMBERSl est branchlets, becoming spiralIy arranged on absent); 43: transition from earlywood to late- thicker ones, simple, scale-hke, on ultimate wood gradual. Tracheids: 44: tracheid pitting branchlets ovate to rhombic, 1.5-3.5 mm x I- in radial walls (predominantly) uniseriate (ear- 1.5 min, on leading branchlets up to 10 min x 4 Iywood only); (45: tracheid pitting in radial mm, apex obtuse to acute, upper margins ini- walls coredominantly) a 2-senate (earlywood nutely toothed, dull pale green. Male cone ter- only)); 56: torus present (pits in earlywood tra- minal on short lateral branchlets, solitary, ob- cheids only); 60: warty layer visible under the long, 3-6 min x 1.5-2 mm, yellowish green light microscope. Axial parenchyma: 72: axial when young, yellowish brown to brown when parenchyma present; (73: axial parenchyma mature; scales 4-8, decussately opposite, pel- diffuse (evenly scattered throughout the entire tate, each bearing 3-5 pollen sacs. Female cone growth increment)); 74: axial parenchyma tan- lateral, sometimes terminal, solitary or gentiany zoriate; 76: transverse end walls grouped, irregularly globose when mature, 15- smooth. Ray composition: 85: end walls of ray 22 mm in diameter, brown or blackish brown, parenchyma cells smooth (unpitted); 87: bon- 3-Toe18)-seeded; scales 4(-6), decussately zorital walls of ray parenchyma cells smooth opposite, woody, oblong, outer surface smooth (unpitted). Cross-field pitting: 93: cross-field to rough. Seeds ovoid, flattened, 5-7 mm long, pits cupressoid; 98: I-3 pits per cross-field blackish brown or black, with 2 wings up to 3 (earlywood only). Ray size: (102: average ray min wide. height very low (, 4 cells)); 103: average ray Other botanical information Widdringto- height medium (5-15 cells); 107: ray width itto comprises 4 species, all in southern Africa. exclusively uniseriate. Widdringtortio whytei has sometimes been in- (P. Baas & I. Hem") cluded in the more widespread Wtddringtonio Growth and development The growth of nodi/10ro (L. ) POWrie (mountain cypress or Widdringtonio whytei in natural stands Is Cape cypress), but it is now considered a sepa- slow. It does not coppice after felling or above- rate species. Widdringtonio nodiftoro is a ground destruction (e. g. by fire or rodents). The multi-stemmed shrub or narrow-crowned small seeds are dlspersed by wind to medium-sized tree up to 25 in tall, with a Stands of about 50 years old in Malawi had a hole diameter up to 50 cm. It is distributed mean hole diameter of 42 cm from Malawi (including Mount Mulanje) Ecology Waddringtonio whyteioccurs scattered through Zimbabwe and Mozambique to the in Afromontane forest at 1800-2550 in alti- Cape Province in South Africa, and occurs at tude, in regions with abundant precipitation, 100-2600 in altitude, predominantly in cool much of it as fog. Nowadays it mainly occurs in and wet mountain fynbos. In contrast to Wtd- fire-protected valleys. It is a pioneer species driftgtonio whytei, it sprouts from the base unable to regenerate under closed canopy after fire and it Is common in fire-prone habi- Therefore, seedlings are either found occasion- tats. Waddringtonto nodtftoro is economicalIy ally on forest edges, or in larger numbers after much less valuable than Widdringtonio whytei; a fire or landslide. Young trees are killed by it is too small to become important as a source fire, but older ones may survive mild fires due of timber, and its wood has a lower density. It to their thick bark is used for hut construction, and probably for Propagation and planting Widdringtonio firewood as well. Extensive plantations of Wid- whyteiis easily propagated from seed driftgtonio nodi/lord have been established, Diseases and pests The aphid Ginoro CM- under the wrong assumption that they were pressi has become a threat in Malawi and Widdringtonio whytet. In Malawi the first elsewhere; it causes local chlorosis and abscis- plantations were established on Zomba plateau SIon of branchlets, and may killyoung plants around 1900, but, as elsewhere, an unintended Harvesting The wood has been heavily ex- mixture of Widdringtonio whytei and Wid- PIOited in Malawibecause of its general utility dringtonio nodiftoro was sown. As the slower Genetic resources Widdringtonio whyteiis growing Widdringtonio whytei was at a disad- acutely threatened with extinction. Important vantage, Widdringtonio nodi/lord became the threats are overexploitation, fire, poor regen- dominantspecies eration, and invasion of its native area by Anatomy Wood-anatomical description JAWA Pinus potulo Schltdl. & Chain. It is classified softwood codes) as endangered in the 2006 TUGN Red list of Growth rings: (40: growth ring boundaries dis- threatened species. It is officially protected and tinct); (41: growth ring boundaries indistinct or licences are available only for the exploitation

. XANTHOCERCIS 589 of dead trees. Illegal felling or killing of trees is Properties The heartwood is beige-brown to known to take place, as the timber is highly brown, often with lighter patches and dark valued and fetches high prices streaks, and distinctly demarcated from the 6- Prospects The wood of Widdringtonio whytei 8 cm wide and paler sapwood. The grain is is of high quality: it is strong, durable and fra- often wavy, texture fine and even. The wood is grant. The species is, however, severely endan- very heavy, with a density of 1020-1230 kg/ms gered, and the exploitation of living trees is at 12% moisture content. The rates of shrink- prohibited. Effective conservation of the re- age during drying are high: from green to oven maining stands is urgently needed. Replanting dry 6.0-7.7% radial and 11.0-13.7% tangential on Mount Mulanje in Malawiis advocated, but The wood is hard. At 12% moisture content, the the growth rate of the tree is slow. Mixed modulus of rupture is 180-202 N/mm2, modulus planting with Waddrirtgtonio nodiftoro should of elasticity 14,610-17,360 N/mm2, compression be avoided. The essential oil from sawdust of parallel to grain 74-91 N/mm2, shear 11.3-12.3 Widdringtonio whytei will not become impor- N/min2, cleavage 36-43 N/min and Chalais- tant in international trade, because the pro- Meudon side hardness 10.9-12.4. duction potential is too low, but it may find The wood is rather difficult to saw, requiring some application in locally manufactured fra- much power and dulling sawteeth rapidly. It is grance products difficult to work, and nailing is almost impos- Major references Chalk, Burtt Davy & SIble without pre-boring. The wood finishes Desch, 1932; Chapman, 1994; Chapola, 1990; well. Staining and polishing give very good Fanon, 2005; Green, Wood & Robinson, 1988; results. The heartwood is durable and resistant Pauw, 1998; Pauw & Linder, 1997; UNEP- to fungi, termites and marine borers. The sap- WCMC, 2006; Williamson, 1955. wood, however, is liable to blue stain and not Other references Chudnoff, 1980; Ciesla, durable. The sawdust irritates mucous mein- 1991; Coates Palgrave, 1983; Conifer Specialist branes and protective measures are needed Group, 1998b; Da Graga Silva, 1983; Dallimore during sawing operations & Jackson, 1966; Foot, 1967; Hemz, 2004; Botany Deciduous small to fairly large tree Lewis, 1960; Marsh, 1966; Mayhead & Ofesi, up to 30 in tall; hole straight and cylindrical, 1989; Oliver, 2006; van Wyk & van Wyk, 1997 branchless for up to 10 in and up to 80 cm in Sources of illustration Fanon, 2005 diameter; bark smooth with many lenticels, Authors M. Brink becoming flaky. Leaves alternate, imparipinnately compound with (5-)7-9(-10) leaflets; petiole and rachis glabrous; leaflets alternate, XANTHOCERCISMADAGASCARIENSIS Baill ovate to oblong-ovate, 3-8 cm x 2-5 cm, rounded shallowly cordate at base, acuminate Protologue Adansonia 9: 294 (1870) at apex, thinly leathery, glabrous. Inflores- Family Papilionaceae (Leguminosae - Papil- cence an axillary or terminal raceme, often ionoideae, Fabaceae) branched and combined into a terminal pani- Origin and geographic distribution Xon- cle, many-flowered. Flowers bisexual, papil- thocercis mudoguscoriensis is endemic to ionaceous; pedicel with 2 small bracteoles; ca- Madagascar, where it mainly occurs in the Iyx broadly cup-shaped, 3-4 min long, truncate northern half. at apex, densely yellowish hairy; corolla Uses The wood is used sometimes as a sub- creamy to pinkish, with oblong standard c. I stitute of the wood of Guoiocum officingIe L cm X 0.5 cm, shortly clawed at base, UPCurved, for luxury furniture, exterior and interior join- wing and keel petals similar, narrowly oblong, ery, luxury flooring, and in boat construction all 5 petals with a yellowish hairy stripe out- It is also used for wood carving, and for making side; stamens 10, fused at extreme base; ovary butcher's blocks, pulleys and luxury decorative superior, shortly stalked, shortly hairy, I- handles of brushes and implements. The wood celled, style short, stigma indistinct. Fruit a is suitable for mine props, vehicle bodies, musi- drupe-like elhpsoid pod 2.5-5(-9) cm x 1.5-2.5 calmstruments, toys and novelties, turnery, cm, slightly constricted between the seeds, draining boards and pattern making. It is glabrous, smooth, indehiscent, I-2(-3)-seeded, popular for poles that are durable in contact with seeds enclosed in a fibrous pulp. Seeds with the ground. The wood is used as firewood kidney-shaped, 1.5-2.5 cm x I-1.5 cm, dark and for making charcoal. The fruit pulp is edi- brown. ble. Xonthocercis mudogoscoriensis can fix atmos-

. 590 TIMBERSl pheric nitrogen by symbiosis with rhizobia XERODERRISSTUHLMANNllCTaub. )Mendonqa Xonthocercis comprises 3 species: one ln Mada- & E. G. Sousa gascar, one in Gabon and one in southern at- Tica. It is related to Angy!ocolyx Protologue Bol. SOC. Brot. , ser. 2, 43: 273 Xonthocercis inodogoscoriensis closely resem- (1968) bles Xonthocercis robiensis Maesen from Ga- Family Papilionaceae (Leguminosae - Papil- bon, which differs in having up to 5 leaflets per ionoideae, Fabaceae) leaf and greyish hairy calyx. Xonthocercis robi- Chromosome number2n=24 errsis is a large tree up to 40 in tall with a cy- Synonyms OStryoderris stuhlmonnii(Taub. ) jindricalbole up to 300 cm in diameter, known Harms (1915). only from a few specimens. Its wood is slightly Vernacular names Wing pod, wing bean lighter in weight than that of Xonthocercis (En). Micala (Po). Mnyinga, mondogondo, inu- incdogoscoriensis, with a density of 950-990 inundu (Sw). kg/ina at 12% moisture content. Xonthocercis Origin and geographic distribution Xero- robiensis is classified as vulnerable in the derris stuhlmonnii is widespread in tropical IUCN Red list of threatened species Africa, from Senegal east to Kenya and south Xonthocercis 20mbesioco (Baker) Dumaz-Ie- to Zimbabwe, Mozambique and northern South Grand (nyala tree) occurs in southern Africa Africa Its wood is occasionally used although it Uses The wood is used for furniture, railway strongly irritates nose and throat during work- sleepers, canoes, handles and utensils. In Tan- ing, but the use of its edible fruits is more jin- zania the wood is locally used to make grain portant. mortars. In Ghana the branches are used for Ecology Xonthocercis mudogoscoriensis oc- making roofs. The wood is suitable for heavy curs in deciduous and evergreen forest up to construction, flooring, vehicle bodies, sporting 400 in altitude. It can be found on sandy soils, goods, Interior trim, joinery, poles and piles, but also on limestone outcrops and basaltic carving, toys and novelties, turnery, veneer soils and plywood. It is also used as firewood and for Genetic resources and breeding Xontho- the production of charcoal cercis ingdogoscorte, 1sts is classified as unlner- The bark contains a fibre, which is sometimes able in the IUCN Red list of threatened spe- used to make brooms. In Zambia the bark is cies. It is felled selectively for its timber and used for tanning hides. The red bark exudate is this makes it liable to genetic erosion, although sometimes used as a dye and for tanning it is fairly widely distributed and occurs within Seeds are eaten in times of food shortage after some protected areas. prolonged cooking, as they are reported to be Prospects Xonthocercis ingdogoscoriensis is poisonous in fresh condition. They are also an interesting timber tree reaching fair dimen- ground to produce a meal. In Tanzania sions and providing excellent-quality timber. Xeroderris stuhlmonniiis considered useful for However, logging in the remaining natural wind breaks. Leaves and fruits are browsed by populations should be done with great care as livestock the species has already become vulnerable. The bark is used in traditional medicine to Tests on planting Xonthocercis ingdogos- treat coughs, colds, rheumatic arthritis, stom- curiensis in Madagascar seem worthwhile to ach-ache, dysentery, eye infections, wounds, determine its potential as a plantation timber and as a purgative. Root decoctions serve as a tree. However, slow growth is likely, coinpara- vermifuge; they are taken to treat elephantia- ble to Xonthocercis robiensis, for which a mean SIS, gonorrhoea, syphilis, dysmenorrhoea, chest annual diameter growth of 4 mm has been re- pains and urinary complaints, and applied corded under natural growing conditions externalIy against ringworm. Pounded roots Major references Bolza & Keating, 1972; are applied to sores. The leaves are used to Derienne, 1998; du Puy at a1. , 2002; Gu6neau, treat colds, coughs, wounds, stomach problems, Bedel & Thie1, 1970-1975; Takahashi, 1978. amenorrhoea, fever and malaria Other references du Puy & Labat, 1998m; Properties The heartwood is cream-coloured Parant, Chichignoud & Rakotova0, 1985; to dark yellow, occasionally with reddish Peltier, 1972; Schatz, 2001; Sprent & Parsons, streaks, and not distinctly demarcated from 2000; van der Maesen, 1997; World Conserva- the sapwood. The grain is straight, rarely in- tion Monitoring Centre, 1998e. terlocked, texture moderately fine and even. Authors R. H. M. J. Lemmens The wood is heavy, with a density of 800-835

. XYi, IA 591 kg/ms at 12% moisture content, and fairly fruits are sometimes inalformed and berry-like, hard. It air dries slowly, with considerable dis- which is caused by galls. Nodulation and nitro- tortion but little checking; kiln drying gives genase activity have been recorded for the best results. Shrinkage rates from green to roots, indicating atmospheric nitrogen fixation 12% moisture content are 3.1% radial and 4.5% Xeroderris comprises a single species. It is re- tangential. Once dry, the wood can be unstable lated to Agonope in service. The surface of the wood can be given Ecology Xeroderris stuhlmonnii occurs in a smooth, lustrous finish. The nailing, screwing deciduous woodland and bushland up to 1650 and gluing properties are good. The wood is in altitude. It grows particularly on well- moderately durable;it is susceptible to powder- drained soils, also on poor sandy soils, and is post beetle attack, but not particularly to ter- drought resistant mite and pinhole borer attack. The wood is ManagementXeroderrisstuhlmonrtitcan be moderately resistant to impregnation. propagated by seeds and suckers. There are The ultimate fibres are 1.5-1.6 mm long and 3500-5000 seeds per kg. The germination rate 20-23 pin wide, with a lumen diameter of 5 pin is up to 70% 2 weeks after sowing. The tree and a cellwallthickness of about 8 prn. Due to tolerates pruning, Dollarding and coppicing the narrow lumen and thick walls, the fibres Genetic resources and breeding Xeroder- are rigid and the wood is not suitable for the ris stuhlmonniiis widespread and locally production of good quality paper. common, especially in East and southern at- Root decoctions have reportedIy violent action rica, and not under threat of genetic erosion on the intestines, causing strong emesis and Prospects Xeroderris stuhlmonnzL Is a inul- may cause loss of sight tipurpose tree that deserves more research Botany Smallto medium-sized tree up to 18(- attention. Although it is widely used in tradi- 27) in tall; hole branchless for up to 12 in, tional medicine, very little is known about its straight and cylindrical, up to 120 cm in di- phytochemistry and pharmacological proper- ameter; bark usually rough, flaking, grey- ties. Detailed information is needed on natural brown, with a red exudate; crown rounded, regeneration and growth rates to recommend with ascending branches; young twigs appropriate methods of sustainable harvesting brownish hairy, older branches thickened, This is needed because the most commonly corky, with distinct leaf scars. Leaves alternate harvested parts, apart from the timber, are and often clustered near the branch tips, jin- bark and roots for medicinal purposes, and panpinnately compound with (5-)6-8 pairs of harvesting these may also knithe tree leaflets; stipules linear-lanceolate, 6-12 mm Major references Arbonnier, 2000; Bolza & long, caducous; leaflets more or less opposite, Keating, 1972; Burki11, 1995; Mbuya at al. , oblong-lanceolate to oblong-ovate, 4-13 cm x 1994; Palmer & Pitman, 1972-1974 2.5-6.5(-9) cm, asymmetricalIy rounded or Other references Asase at a1. , 2005; Bryce, cordate at base, usually rounded at apex, mar- 1967; Coates Palgrave, 1983; Gelfand at al. , gins slightly recurved, leathery, golden or SII- 1985; Huxham at a1. , 1998; Kokwar0, 1993; very silky hairy when young, later glabrescent. Neuwinger, 2000; Taita, 2000; Tropical Prod- Inflorescence an axillary panicle 7-18(-22) cm ucts Institute, undated; van Wyk & van Wyk, long, crowded at the ends of twigs, with spread- 1997 ing branches, rusty hairy, many-flowered. Authors R. H. M. J. Lemmens Flowers bisexual, papilionaceous; pedice1 2-5 mm long; calyx broadly campanulate, 4-5 mm long, hairy; corolla c. 1.5 cm long, white or XYLIAEVANSilHutch. greenish white, standard circular to oblate, with short claw, wings and keel about equally Protologue Bun. Msc. Inform. Kew 1908: 258 long; stamens 10, 9 fused and I free; ovary (1908) superior, sessile, I-ceUed, style curved, stigma Family Mimosaceae (Leguminosae - Mimo- small. Fruit a linear-oblong pod 9-Ise30) cm x soldeae) 2-5 cm, flattened, with a wing at both sides, Chromosome number2n=24 pale brown, reticulately veined, indehiscent, I- Origin and geographic distribution Xy!to 3(-5)-seeded. Seeds kidney-shaped, c. 12 min euonsiioccursfrom Sierra Leone to Ghana long, smooth, red-brown Uses The wood is used for local construction. The tree often flowers at the end of the dry The twigs are used as chewing sticks, and a season, shortly before new leaves develop. The decoction of leafy twigs is administered as a 592 TIMBERSl cholagogue and tonic. The leaves and ash from Young Xylio euonsiitrees have an annual di- pods are used as a substitute for soap. Roasted ameter increment of 0.6-1.4 cm. Seedlings de- seeds are reportedIy edible. A vegetable salt velop ectotrophic mycorrhizae. The trees are has been obtained from wood ash by Ieaching. often deciduous for a short period. Flowers are Properties The heartwood is reddish brown produced towards the end of the dry season. with darker streaks and distinctly demarcated The seeds are dispersed by the explosiveIy from the pale yellow sapwood. The grain is opening pods at the end of the dry season interlocked; the texture fine. The wood is inod- Ecology Xylio euunsii occurs in evergreen erately heavy, with a density of about 770 forest, semi-deciduous forest and gallery forest. kg/ms at 12% moisture content, and hard. It It is often found on well-drained alluvial soils dries quite fast and well. The recorded worka- and on hillsides with deep soils hinty of the wood varies from fairly well to dif- Management Large trees usuaUy occur scat- ficult. It is resistant against both fungi and tered in the forest, but are locally abundant, Insects. e. g. in Sierra Leone and Ghana. In some forest Botany Medium-sized to fairly large tree up areas of Sierra Leone an average density of 2.7 to 35 in tall; hole often fluted and bent, up to trees of over 70 cm bole diameter per ha has 160 cm in diameter, with large, humped but- been recorded. Regeneration is in small to me- tresses; bark brown, roughly scaly. Leaves al- dium-sized forest gaps, but seedlings are usu- ternate, clustered at the ends of twigs, hipin- ally not abundant. However, the germination nately compound with a single pair of pinnae; rate of seeds in the nursery is fair. Germina- stipules linear, persistent; petiole 1.5-5 cm tion starts 4-10 days after sowing. The 1000- long, shortly hairy, with large gland at apex on seed weightis about 400 g. upper side; pinna axis 10-35 cm long; leaflets Genetic resources and breeding Xylio in 9-20 pairs per pinna, opposite, oblong- euonsii does not seem to be in Immediate dan- lanceolate to oblong-elliptical, up to 9 cm x 2 ger of genetic erosion. Although it is restricted cm, rounded at base, acuminate at apex, to the forest zone of West Africa, it is wide- shortly hairy below. Inflorescence an axillary spread there, and locally common head c. 2 cm in diameter, many-flowered; pe- Prospects Little is known about Xylio euon- duncle 3-8 cm long. Flowers bisexual, regular, sit, and more research, especially on propaga- 5-merous, small, sessile, hairy; calyx cylindri- tion and growth rates, is needed to evaluate its cal, c. 3 min long, shortly toothed; petals free, possible role in sustainably managed natural linear-oblong, c. 4 min long, brownish yellow; production forest in West Africa stamens 10, free, c. 8 mm long, with glands at Major references Burki11, 1995; Holingren apex; ovary superior, c. I mm long, long-hairy, at a1. , 2004; Saville & Fox, 1967. I-celled, style short. Fruit a narrowly oblong Other references Aubr6ville, 1959c; de in pod up to 20 cm x 5 cm, flattened, long- Mensbruge, 1966; Dudek, F6rster & mis- attenuate at base, obtuse at apex, slightly senbauer, 1981; du Puy at a1. , 2002; Haw- curved, woody, brown, 2-valved, 4-9-seeded thorne, 1995; ITvine, 1961; Nguyen Ba, 1998; Seeds obovoid-ellipsoid, c. 2 cm x 1.5 cm, nat- Normand & Paquis, 1976 tened, glossy. Seedling with epigeal germina- Authors R. H. M. J. Lemmens tion. Xylio comprises 9 species, 6 of which occur in continental Africa, 2 in Madagascar and I in ZANTHOXYLUMDAWl(Verd. ) P. G. Waterinari tropical Asia. It is related to Golpocolyx, which differs in its spike-like inflorescences Protologue Taxon 24: 363 (1975). Xyliu xylocorpo (Roxb. ) Taub. from tropical Family Rutaceae Asia Is an important timber tree in Myanmar Synonyms Fuguro douyiVerd. (1919) and India, and has occasionally been planted in Vernacular names kilobwood, forest knob- tropical Africa. The wood is heavy, hard and wood, knobthorn (En). durable and used for heavy construction. The Origin and geographic distribution Zori- wood ofXy!to hoffmo, !nit(Vatke) Drake is used thoxylum douyi occurs in Zimbabwe, eastern for furniture in northern Madagascar; a decoc- South Africa and Swaziland. tion of the pods is taken as a tonic. Xylio hoff- Uses The wood is used for handles, walking monniiis a tree up to 25 in tall, with a bole up sticks and fishing rods. It is suitable for heavy to 40 cm in diameter, and is locally common in construction, heavy flooring, Joinery, interior tallforest on limestone soils trim, vehicle bodies, furniture, cabinet work,

. ZANTHOXYLUM 593 mine props, ladders, sporting goods, agricul- lanceolate or narrowly elliptical, 2-7(-10) cm x turalimplements, toys, novelties, musical in- I-3 cm, curieate to obtuse at base, shortly struments, boxes, crates and turnery acuminate and slightly notched at apex, mar- The stem bark is used to treat cough, cold, gin finely toothed, glabrous, with glandular boils, pleurisy, toothache and snakebites. The dots confined to the margin, pinnately veined prickles are applied to infected wounds, the with numerous lateral veins. Inflorescence a leaves to treat chest pain and as a poultice to terminal panicle up to 6 cm long, with flowers heal sores, and the roots to treat mouth ulcers in clusters. Flowers unisexual, regular, 4- and sore throat, and as a tonlc and aphrodisiac merous, small; pedice1 I-1.5 min long; sepals The prickle-bearing protuberances on the hole nearly free, c. 0.5 mm long; petals elliptical, c have been used by children as toys, and have 2.5 min long, greenlsh yellow; male flowers also been made into pines with 4 stamens, ovary rudimentary; female Properties The heartwood is pale greyish flowers with superior, globose ovary and short brown, often with greenish markings, fairly style. Fruit a globose follicle c. 5 min in diame- distinctly demarcated from the greyish sap- ter, glandular pitted, dehiscent, I'Seeded. Seed wood. The texture is fine and even. Growth globose, c. 3 mm in diameter, black and shiny rings are distinct Zanthoxylum douyiflowers from October to The wood is heavy, with a density of 830-930 January and fruits ripen about 3 monthslater. kg/in3 at 12% moisture content It air dries Zanthoxylum is pantropical and comprises fairly rapidly in small dimensions, but larger about 200 species, with tropical milerICa being boards are liable to surface checking. The rates richest in specles Mainland Africa harbours of shrinkage are moderate, from green to oven about 35 species, whereas about 5 species are dry 4.1% radial and 8.4% tangential. endemic to Madagascar The wood is strong and elastic. At 12% ino1s- Ecology Zorithoxylum douy, occurs in forest ture content, the modulus of rupture is 128- and forest patches, from coastal regions to 140 N/mm2 modulus of elasticity 15,700 mountainous areas; it is locally common In N/mm2, compression parallel to grain 55-57 montane forest. N/mm2 shear 15-17 N/min2, Janka side hard- Genetic resources and breeding The con- ness 9210-10,450 N and Janka end hardness servation status of Zorithoxylum douyi in 9790-11,260 N South Africa is indeterminate, but in Zim- The wood is fairly easy to saw and work in babwe it is considered endangered spite of its hardness. The polishing, gluing and Prospects The information on Zorithoxylum bending properties are satisfactory. The wood douyiis too limited to Judge Its prospects as a is only moderately durable and susceptible to commercial timber tree under sustainable ex- Lyetus attack. The heartwood is resistant to PIOitation management. However, stands of impregnation by preservatives large trees are probably too scarce, and an in- The stem bark yielded 5 benzo[c]phenanthridine ventory is needed. alkaloids. One of these, chelerythrine, is well The medicinal properties deserve more atten- known for its antimicrobial and anti- tion because of the claimed antimicrobial ac- inflammatory properties. Crude stem extracts tivities, which have been confirmed by phar- showed pronounced antibacterial activity macologicalresearch. against ESCherichio coli, Shige!10 spp. , Solmo- Major references Bolza & Keating, 1972; itel!o typhi and Streptococcus pyogenes, but no Coates Palgrave, 1983; Palmer & Pitman, activity against Stophy!ococcus oureus. 1972-}974; Takahashi, 1978; Tarus at al. , Botany Dinedous, medium-sized tree up to 30 2006 in tan; hole usually straight and cylindrical, up Other references Grace at a1. , 2002a; Kel- to 60 cm in diameter, with conspicuous woody, manson, J6. ger & van Staden, 2000; Neu- prickle-bearing protuberances, sometimes winger, 2000; Obi at a1. , 2002; van Vuuren, slightly fluted at base; bark pale grey in Banks & Stohr, 1978; van Wyk & van Wyk, younger trees to dark brown in older ones; 1997 twigs glabrous, armed with up to 5 mm long Authors R. H. M. J. Lemmens prickles. Leaves alternate, imparipinnately compound with (5-)7-13 leaflets, up to 30 cm long; stipules absent; rachis channelled above, often with small prickles; leaflets opposite, sessile, slightly asymmetricalIy oblong to 594 TIMBERSl

ZANTHOXYLUM GILLETli(De Wild. ) Terns including kidney complaints and gonor- P. G. Waterman rhoea, as a vermifuge and as an enema against severe diarrhoea. The bark is applied exter- Protologue Taxon 24: 363 (1975). nany to treat cough, colds, skin complaints and Family Rutaceae smallpox. It is also used as fish poison and Chromosome number2n=64 arrow poison. The leaves are used to treat Synonyms Fuguro inocrophy!IQ (011v. ) Engl heart complaints and snake bites, whereas a (1896), Foguro tessmonniiEng1. (1911), Fogoru leaf decoction is taken to treat cough, gonor- Qinoniensts Eng1. (1917), Zoritho, cy!urn tessmon- rhoea and schistosomiasis, and a leaf macera- nit(Engl. ) J. FAyafor (1984). tion to treat diarrhoea and gastritis. Leaf sap Vernacular names African satinwood, white is applied externalIy against an enlarged African mahogany (En). 010n dur, citronnier spleen d'Mrique, faux citronnier (Fr) The stem protuberances are used as plugs and Origin and geographic distribution Zori- the roots are used as chewing sticks to clean tho"y!urn gilletiiis widespread, occurring from the teeth. Young shoots are added to sauces for Guinea and Sierra Leone east to Kenya and flavouring and in Kenya leaves are added to south to northern Angola, Zimbabwe and Mo- tea for this purpose. Pollen and nectar are col- zambique. It is fairly commonly planted within Iected from the flowers by honey bees. The its distribution area, often as an individual seeds yield an edible oil. Zorithoxylum gilletiiis tree, but sometimes also in pure stands occasionally planted as an ornamental shade Uses The wood, often traded as 'o10nvogo' or tree. o10n', is used for construction in house build- Production and international trade The ing, for flooring, Joinery, interior trim, panel- timber of Zorithoxylum gilletii has little impor- ling, doors, shipbuilding, vehicle bodies, furni- tance on the international market and most of ture, cabinet work, mine props, railway sleep- the trade is for local use. Trade statistics are ers, handles, ladders, sporting goods, agricul- not available, but the officially registered tim- tural implements, drums, toys, novelties, her extraction from Kakamega Forest in Kenya boxes, crates, turnery, veneer and plywood. declined from 645 in31year in the 1930s to less The wood is also used as firewood and for char- than 100 ing in 2000 coalproduction Properties The heartwood is pale yellow to Zorithoxylum gilletiiis important in traditional bright yellow or yellowish brown, darkening medicine. The bark of stem and roots is coin- upon exposure, and indistinctly demarcated monly used as an analgesic, especially to treat from the narrow sapwood. The grain is inter- burns, rheumatism, headache, stomach-ache, locked, texture fine to moderately coarse. Quar- toothache and pain after childbirth. The bark tercut surfaces show a stripe figure and back- is also taken against colic and fever and is con- sawn surfaces occasionally have a fiddleback sidered to have aphrodisiac properties. Bark figure. Freshly cut wood has a sweet scent. The decoctions are taken agalnst urogenital prob- wood has a silky lustre The wood is moderately heavy to heavy, with a density of (550-)720-1040 kg/ms at 12% ino1s- ture content. It air dries fairly rapidly with little degrade, although cracks may develop in , . boards. Boards of 2.5 cm thick can be air dried in 6 weeks, boards of 5 cm thick in 3 months Kiln drying usually does not cause problems. However, the rates of shrinkage are moderate ^ to high, from green to oven dry 4.3-6.5% radial ,^ and 8.7-10.2% tangential.

L. . The wood is strong and hard. At 12% moisture content, the modulus of rupture is (86-)1/4-

.. 266 N/mm2, modulus of elasticity 8500-18,400 N/mm2, compression parallel to grain (48-)58- Q 104 N/min2, compression perpendicular to grain 7 N/mm2, shear 10 N/min2 cleavage 16- 25 N/min, Janka side hardness 3650-7790 N, Zorithoxylum gilletii - wild Janka end hardness 5330-9120 N and Chalais-

. ZANTHOXYLUM 595

Meudon side hardness 5.2-13. The wood usually saws moderately well, but is rather difficult to work. Reduction of cutting angles to 15' gives best results in planing. The wood can be finished to a smooth and lustrous surface, but splitting and a rough finish may occur after machine mortising and boring; good I~~' mouldings are possible when sharp cutters are used. The nailing properties are good, but gluing properties are variable. The steam-bending properties are excellent. The wood is only inod- erately durable and often susceptible to ter- 3 \.,., mite, Lyetus and marine borer attacks. The heartwood is moderately resistant to Impreg- 4 nation by preservatives, the sapwood permeable The major volatile constituents Isolated from $ the bark are the sesqulterpenes germacrene B, Telemene and germacrone. The bark contains isoquinoline alkaloids; the root bark contains larger amounts than the stem bark. The qua- ternary herizophenanthridine nitidine and the isobutylamide fagaramide are major constitu- 'I ~-~ -'~ ;--. - .* ents. The first compound lowered blood pres- 2 ',*' .,***;*" sure in tests with rabbits and showed cytotoxlc activity in tumour celllines, whereas the latter compound and some other ISObutylamides Zorithoxylum gilletii - I, 60se of 601e, ' 2, led/; 3, showed insecticidal and molluscicidal activi- legf base, 4, inIructescence ties. The acridone alkaloid xanthoxohne ISO- Redrown und udopted by Achingd Sotiri Nur- lated from the bark exhibited potent an- human tifeedant activity against larvae of Spodoptero spp. Chelerythrine is another alkaloid isolated to 90(-150) cm in diameter, with many woody, from the bark; it showed analgesic effect and prickle-bearing protuberances up to 3 cm long antibacterial, antifungal and arithelmintic ac- but old trees often lacking these, often with tivities. Skimmianine, a furoquinoline alkaloid indistinct buttresses at base; outer bark grey to also present in the bark, showed sedative, hy- greyish brown, smooth to slightly rough, inner pothermic and antidiuretic effects in rats and bark granular, yellowish brown, often inottled mice The heartwood contains coumarine de- with orange; crown spreading; branches gla- rivatives such as 6.7-dimethoxycoumarine; this brous, armed with conical prickles up to 8 mm compound has anti-inflammatory, analgesic, long. Leaves alternate, clustered at ends of choleretic, hypotensive and tranquilizing prop- branches, imparipinnately compound with 13- erties. Saponins are present in bark and 27(-51) leaflets, up to 120(-150) cm long; stip- leaves. Extracts of root bark and stem bark ules absent; rachis glabrous, sometimes with showed considerable antimalarial activity in prickles; petiolules up to I cm long; leaflets vitro using multi-drug resistant strains of alternate to nearly opposite, elliptical-oblong, Pigsmodium 101ctporum. Leaves showed good (8-)14-30 cm x 3.5-10 cm, curieate to rounded results in the treatment of sickle-cell anaemia and asymmetrical at base, shortly acuminate patients, probably due to the presence of p- to obtuse at apex, margin entire or sometimes hydroxybenzoic acid and similar benzoic acid slightly toothed, leathery, glabrous, with many derrvatives. minute glandular dots, pinnately veined wTth The leaves yield a volatile oil, with an ocLmene 8-18 pairs of lateral veins. Inflorescence a ter- derivative as major constituent (about 43%) minal or axillary pyramidal panicle 20-35 cm and additionany or-pinene and or-phellandrene long, many-flowered with flowers in clusters. Description Deciduous small to fairly large Flowers unisexual, regular, 5-meTous, small, tree up to 35 in tall; hole branchless for up to nearly sessile; sepals united halfway, ovate to 15(-25) in, usually straight and cylindrical, up circular, 05-I min long; petals obovate to fid-

. 596 TIMBERSl

dle-shaped, I-2.5 min long, white, turning prismatic crystals in chambered axial paren- brown; male flowers with 5 stamens varying in chyma cells. length, disk dome-shaped and lobed, ovary (N. P. Monel, P. Derienne & E. A. Wheeler) rudimentary; female flowers with superior, Growth and development Zorithoxylum ovoid ovary I-1.5 mm long, shortly stiped, gilletti grows fairly rapidly. Seedlings may stamens rudimentary. Fruit a globose follicle reach a height of 40-70 cm after the first year, 3.5-6 mm in diameter, reddish, glandular pit- and annual growth rates are 35-50 cm. In C6te ted, dehiscent, I(-2)-seeded. Seed globose, 2.5- d'TVoire trees with a bole diameter of 50 cm 3.5 mm in diameter, black and shiny. Seedling have been found in secondary forest of 40 years with epigeal germination; hypocoty1 3-4 cm old. Trees may start flowering when 10 years long; cotyledons oblong, I-1.5 cm long, leafy; old. Flowering is irregular, probably due to first leaves simple variable climatic conditions. Trees flower for Other botanical information Zorithoxy!urn about 2 months. Fruits ripen about 3 months Is pantropical and comprises about 200 species, after flowering. In Kenya the seeds of most with tropical America being richest in species Zanthoxylum gilletii trees ripen in the rainy Mainland Africa harbours about 35 species, season, which is advantageous for germination whereas about 5 species are endemic to Mada- The seeds are probably mainly dispersed by gascar birds and monkeys. Trees may flower and fruit The wood of Zorith0, o. 1um beequetii(G. CG. Gnuert) Once a year or once In every two years P. G. Waterinari (synonym: Fogoro beeqt, etii Ecology Zanthoxylum gilletii occurs in ever- G. CG. Gilbert) has been used in Rwanda for green rainforest, in East Africa up to 2400 in construction;it is reported as hard and durable altitude. The mean annual rainfall in its area The wood of Zoritho, ;}^Ium renieri(G. CG. Gitbert) of distribution is 1200-2400 min. Zorithoxylum P. G. Waterman (synonym: Fogoro renier! gilletii usually occurs scattered in the forest. In G. C. CGIlbert), occurring in eastern DR Congo West and Central Africa it is a pioneer species andRwanda, has been used for making canoes that is most common in secondary forest. Seed- Anatomy Wood-anatomical description (IAWA lings can be common in burnt forest. Zorithoxy- hardwood codes) Ium gilletiiprefers well-drained soils Growth rings: a growth ring boundaries iridis- Propagation and planting Seedlings are tinct or absent. Vessels: 5: wood diffuse-porous; light-demanding and natural regeneration may 13: simple perforation plates; 22: intervessel be abundant in large gaps in the forest and in pits alternate; (24: intervessel pits minute (S 4 regrowth of old farmland. In Liberia seedlings Urn)); 25: intervessel pits small(4-7 pin); 30 are commonly present in such localities, but vessel-ray pits with distinct borders; similar to reportedIy never gregarious. Fruits should be Jntervessel pits in size and shape throughout collected from the tree before they open but the ray cell; 42: mean tangential diameter of when already reddish brown. They should be vessellumina 100-200 pin; (43: mean tangen- dried in the sun for I-2 days, and subsequently tial diameter of vessellumina ;^ 200 F1m); 46: S the seed can be shaken out. The 1000-seed 5 vessels per square minimetre; (47: 5-20 ves- weight is 15-35 g. Germination starts 3 weeks sels per square minimetre); 58: gums and other after sowing. Seeds are recalcitrant and the deposits in heartwood vessels. Tracheids and germination rate is often low. Germination fibres: 61: fibres with simple to minutely bor- rates of 20-50% have been reported, but in dered pits; 66: non-septate fibres present; 69 western Kenya rates of up to 80% in 75-120 fibres thin- to thick-walled. Axial parenchyma days. The oily and hard seed coat contributes 79: axial parenchyma vasicentric; 80: axial to the often poor germination. Washing seeds parenchyma allform; 81: axial parenchyma thoroughly with a soap solution improves the lozenge-allform; (83: axial parenchyma conflu- germination rate and reduces the germination ent); 92: four (3-4) cells per parenchyma period significantly. In West Africa it has been strand; 93: eight (5-8) cells per parenchyma recorded that germination is rapid and that strand. Rays: 97: ray width I-3 cells; 104: all viahinty of the seed is short. The seed should ray cells procumbent; (106: body ray cells pro- be kept in the shade, and can be stored for up cumbent with one row of upright and/or square to 2 months. To prevent insect attacks, ash marginal cells); 1/5: 4-12 rays per mm. Secre- should be added. Wildlings are commonly used tory elements and cambial variants: 131: inter- for planting because of the erratic germination cellular canals of traumatic origin. Mineral Seedlings are pricked out into tubes at 5-12 inclusions: 136: prismatic crystals present; 142 days after germination, or 3-4 seeds are sown

. ZANTHOXYLUM 597 directly in tubes. The seedlings should be The medicinal properties deserve more atten- grown in the shade and slowly hardened. They tion. Several of the claimed activities in tradi- are ready for transplanting after 5-7 months. tional medicine have been confirmed by phar- The usual spacing is 3.5-5 in x 3.5-5 in, result- macological research, and a next step might be ing in 400-800 sternsftia the development of safe and standardized Management Zorithoxylum gilletii trees plant-based drugs. often occur scattered in the forest in low densi- Major references BOLza & Keating, 1972; ties; for south-western Cameroon an average Burki11, 1997; Katende, Birnie & Tengnas, exploitable timber volume of less than 0.1 1995; Kokwar0, 1982; Latham, 2004; Maundu matha has been recorded. Planting should be & Tengnds, 2005; Neuwinger, 1996; Phong- done at the onset of the rainy season in areas phaew, 2003; Takahashi, 1978; World Agrofor- where irrigation is not possible. Planting holes estry Centre, undated are 30-45 cm in diameter and 45-60 cm deep Other references Ako6gninou, van der In Kenya phosphate fertilizers are applied, and Burg & van der Maesen, 2006; Beentje, 1994; manure is sometimes also applied. At the usual Fouarge & G6rard, 1964; GIIbert, 1958a; Haw- spacing thinning is not needed, but at spacings thorne, 1995; Hawthorne & Jongkind, 2006; of less than 3.5 in x 3.5 in first and second InsideWood, undated; Kubo at a1. , 1984; Le- thinnings at 8 years and 14 years, respectively, touzey, 1963b; Mendonqa, 1963; Neuwinger, are needed. The tree can be managed by COD- 2000; 0guntimein, EI A1fy & EISOhly, 1985; PICing Pauwels, 1993; Raponda-Walker & Sillans, Diseases and pests Damping-offdisease can 1961; Ruberto & Tnngali, 1998; Stepel, Poorter be a serious problem in the nursery, especially & Hawthorne, 2004; Tringali at a1. , 2001; when the soilis poorly drained. Defoliators are Vivien & Faure, 1985; Voorhoeve, 1979; Witks common, but usually not problematic. Borers & Issemb6, 2000; Zirihi at a1. , 2005 eating the terminal bud resulting in forked Sources of illustration Hawthorne & stems have been recorded in Guinea. Seeds Jongkind, 2006; Voorhoeve, 1979; Wilks & Is- maybe commonly infested with insectlarvae semb6, 2000. Harvesting Logging for timber Is usually Authors MM. Okeyo done during the dry season when the forests where Zorithoxylum gilletii occurs can be en- tered more easily. Where demand for the bark ZANTHOXYLUMHEITZll(Aubr6v. & Fellegr. ) exists, logs are debarked after harvesting P. G. Waterman Handling after harvest The wood is susceptible to stain and logs should be removed Protologue Taxon 24: 363 (1975) from the forest immediately after felling or be Family Rutaceae treated with an anti-stain solution Synonyms Fuguro heitziiAubr6v. & Penegr Genetic resources Although Zorithoxylum (1950), Fogor0 67ieyiVermoesen ex G. C. C. GIIbert gilletiiis widespread, there Is some concern (1958) about genetic erosion. Wood joggers target big Vernacular names 010ntendre (ET). straight trees and these have become scarce In Origin and geographic distribution Zori- many regions. In forests in Kenya regeneration thoxy!urn heittit occurs from southern Garner- is often poor due to grazing and heavy pressure o0n and the Central African Republic to Gabon by neighbouring communities and Bas-Congo province of DR Congo. Prospects Zorithoxy!urn gilletii may have Uses The wood, often traded as 'o10n' or good prospects as a plantation timber tree 'light o10n', is suitable for light construction, However, more research is needed on propaga- light flooring, joinery, interior trim, moulding, tion to overcome the often poor seed germina- shipbuilding, furniture, cabinet work, toys, tion and to develop proper techniques for vege- novelties, boxes, crates, turnery, veneer, ply- tative propagation including in-vitro propaga- wood, hardboard and particle board. It is used tion. Suitable methods of management of natu- IOCaUy for canoes, drums and beehlves. It is ral forest where Zorithoxylum gilletiiis an Im- sometimes also used asfirewood portant constituent should also be lnvestigated Zanthoxylum heitzii is used in traditional Abundant natural regeneration has been re- medicine. In Garnero0n scrapings from the corded in logged-over forest, which makes It a stem bark are applied externalIy to treat ina- suitable candidate for sustainable timber ex- Iana, and in Congo to soothe toothache. In Ga- PIOitation bon the bark is applied externalIy in lotions to

. 598 TIMBERSl

Quartercut surfaces show a slight ribbon-like figure. Freshly cut wood has a slight, pleasant scent. The wood is lustrous The wood is medium-weight, with a density of

. 450-560 kg/ina at 12% moisture content. It air dries rapidly with slight risk of distortion and checking. The rates of shrinkage are moderate, from green to oven dry 3.7-3.9% radial and 5.7-6.1% tangential. Once dry, it is moderately ,^ stable In service

L. , At 12% moisture content, the modulus of rupture is 72-122 N/mm2, modulus of elasticity

. 10,000-12,400 N/min2, compression parallel to grain 35-51 N/mm2, shear 6 N/min2, cleavage o 7-12 N/mm and Chalais-Meudon side hardness 1.5-2.9 The wood saws and works well with ordinary Zanthoxylum heitzii - wild machine and hand tools. Planing may cause some difficulties due to interlocked grain. The treat rheumatism and stiffness, and a macera- moulding, polishing, gluing, painting and var- tion of young twigs in lemonjuice to treat heart nishing properties are all satisfactory. The complaints. In Garnero0n the bark is used to screwing and nailing properties are good and treat gonorrhoea, abscesses and painful joints the wood holds screws well, but nails are easy and male sexual impotence. In Congo the bark to draw. It produces good veneer by slicing and Is used as an analgesic. It serves as a fish poi- rotary cutting. The wood is not durable and son. The tree is sometimes retained by farmers susceptible to fungal, termite, Lyetus and ina- after clearing the forest for agricultural land, rine borer attacks. The heartwood is resistant to serve as a shade tree for cultivation of crops to Impregnation with preservatives, the sap- such as coffee and cocoa wood moderately resistant. The sawdust may Production and international trade The be irritant; allergic contact dermatitis has been annual production of Zanthoxylum heitsit tim- recorded in workers her in Gabon in 1994-1995 has been estimated The wood contains: cellulose 41-47%, pento- at 5700 in3. The timber of Zorithoxylum heitsit sans 10-14%, 11gnin 30-31% and ash 0.1-0.2% has little Importance on the international mar- Solubilities are 1.8-2.4% in hot water, 3.2- ket and is mostly traded for local use. Few 5.8% in alcohol-benzene and 11.1-12.7% in I% trade statistics are available because the wood NaOH. The wood fibres are 1.17 min long and of Zorith0, o. 1um gilletii roe Wild. ) P. G. Waterinari 30 prn wide, with a lumen diameter of 21 pin Is sometimes sold as 'o10n' too. annual o10n log and a cell wanthickness of 8 FLm. Good pulps exports during the 1960s were 5000-6000 in3 can be obtained from the wood using the kraft from Equatorial Guinea and 1300-3000 in31year process and other chemical pulping processes from Gabon, whereas only very small volumes The secondary alkaloid flindersine has been were exported from Cameroon and Congo. In extracted from the heartwood; the bark yielded 1997-1998 Equatorial Guinea exported about several 11gnans, the triterpenes Iupeol and Iu- 1900 ms/year. According to ATIBT statistics, peone, and the alkaloids skimmianine and 6- Gabon exported 310 ina in 2001, 499 ms in methylnitidine. 2002, 1071 ms in 2003 and 886 ina in 2004 Adulterations and substitutes The wood Sawnwood exports are insignificant. of Zanthoxylum gilletii is often sold as 'o10n', The bark is sold on local markets for medicinal although it has a higher density and it is purposes, e. g. in Yaound6 (Cameroon), where harder. in 2002 the price was 5-8 XAF (CFA Central Description Medium-sized to fairly large Franc) per g tree up to 35 in tall; hole branchless for up to Properties The heartwood is pale yellow to 20 in, usually straight and cylindrical, up to greenish yellow and indistinctly demarcated 150 cm in diameter, with many woody, prickle- from the pale yellow, I-2 cm wide sapwood. bearing protuberances up to 9 cm long but old The grain Is usually interlocked, sometimes trees often lacking these, usually without but- straight, texture fine to moderately coarse tresses but often slightly thickened at base;

. ZANTHOXYLUM 599

ovary rudimentary; female flowers with sripe- it-.,., * \, nor, globose ovary I-1.5 mm long, stamens * rudimentary. Fruit a globose follicle c. 4 min in 6 diameter, glandular pitted, dehiscent, I'Seeded.

I Seed globose, c. 2.5 min in diameter, black and shiny. Other botanical information Zorithoxylum is pantropical and comprises about 200 species, with tropical America being richest in species Mainland Africa harbours about 35 species, whereas about 5 species are endemic to Mada- ^\ gascar Zorithoxylum hattziiis often confused 4 5 with Zoritho"ylum gilletii, which produces a heavier and harder timber and also differs in less numerous and larger leaflets with often

. ,,~ entire margins. , ,,. Anatomy Wood-anatontical description OAWA , up* hardwood codes): Growth rings: 2: growth ring boundaries iridis-

'*,,, ' tinct or absent. Vessels: 5: wood diffuse-porous; 13: simple perforation plates; 22: intervessel pits alternate; (23: shape of alternate plts po- I. _ .-~. , , Iygona1); 25: intervessel pits small (4-7 prn); 30: vessel-ray pits with distinct borders; simi- 2 3 '., 11' . If' Iar to intervessel pits in size and shape '. *~-~ throughout the ray cell; 42: mean tangential Zorithoxy!urn heitzii - I, 60se of 601e, ' 2, jeer/; 3, diameter of vessellumina 100-200 prn; 47: 5- leaflet, 4, mole inflorescence, . 5, fernole milores- 20 vessels per square minimetre; (48: 20-40 eeriee, 6, fruit vessels per square minimetre); 58: gums and Redrawn ond oddpted by Achingd Sotiri Nur- other deposits in heartwood vessels. Tracheids hornon and fibres: 61: fibres with simple to minutely bordered pits; 66: non-septate fibres present; outer bark grey to greenish grey, smooth to 69: fibres thin- to thick-walled. Axial paren- slightly fissured and scaly in old trees, lriner chyma: 78: axial parenchyma scanty paratra- bark granular to fibrous, yellowish brown, of- chea1; (79: axial parenchyma vasicentric); (89: ten inottled with orange; crown large, with axial parenchyma in marginal or ln seemingly ascending, strongly branched branches; marginal bands);(91: two ceUs per parenchyma branches glabrous, armed with conlcalprickles. strand); 92: four (3-4) cells per parenchyma Leaves alternate, clustered at ends of strand. Rays: (97: "ay width I-3 cells); (98: branches, imparipinnately compound with 25- larger rays commonly 4- to 10'seriate); 104: all 51 leaflets, up to 100 cm long; stipules absent; ray cells procumbent; (106: body ray cells pro- rachis glabrous, sometimes with smallprickles; cumbent with one row of upright and/or square leaflets nearly opposite, sessile, oblong to ob- marginal cells); 115: 4-12 rays per min. Secre- long-lanceolate, 4-15(-20) cm x 2-5 cm, tory elements and cambial variants: (131: in- rounded to slightly cordate and asymmetrical tercellular canals of traumatic origin). Mineral at base, shortly acuminate at apex, margin inclusions: 136: prismatic crystals present; 142: slightly toothed with rounded teeth, leathery, prismatic crystals in chambered axial paren- glabrous, with scattered minute glandular chyma cells dots, pinnately veined with 10-20 pairs of Iat- (N. P. Monel, P. 06tienne & E. A. Wheeler) eral veins. Inflorescence a terminal or axillary Growth and development Zorithoxy!urn panicle up to 40 cm long, many-flowered with heitzii grows rapidly. The seeds are probably flowers in clusters. Flowers unisexual, regular, dispersed by birds. 5-merous, small, sessile; sepals united for most Ecology Zorithoxylum heitzii occurs in ever- of their length, 0.5-I mm long; petals oblanceo- green and semi-deciduous forest, up to 1200 in late, c. 2 min long, white, turning brown; male altitude. It is a pioneer species that is most flowers with 5 stamens, disk conical, thick, common in secondary forest. It prefers well- 600 TIMBERSl

drained soils Other references Adjanohoun at al. (Edi- Propagation and planting Seedlings are tors), 1988; ATIBT, 1986; ATIBT, 2005; Betti, light-demanding and natural regeneration may 2001; Betti, 2002; Betti, 2003; Bongui at al. , be abundant in larger gaps in the forest. Seeds 2005; Cond6-Salazar, 1987; Foahom, 2002; may germinate 25-30 days after sowing. Seeds Gassita at al. (Editors), 1982; GIIbert, 1958a; that have been soaked in water for longer pen- InsideWood, undated; Letouzey, 1963a; Le- ods do not germinate. touzey, 1963b; Ngavoura, 1990; Ngouela, Management Mature Zanthoxylum heitsit Tsamo & Connolly, 1994; Nkeoua & trees usually occur scattered in the forest. For Boundzanga, 1999; Raponda-Walker & Sillans, southern GarnerDon average exploitable timber 1961; Tailfer, 1989; Vivien & Faure, 1985; volumes of less than 0.1 mama to 0.8 matha Wilks & TSSemb6, 2000 have been recorded, and for Gabon an average Sources of illustration Letouzey, 1963a; timber volume of 0.3 mama Wilks & TSSemb6, 2000 Diseases and pests The wood-boring beetle Authors R. B. Jiofack Tafokou Apote monochus and the hemipterous Lendo punctoto have been recorded as pests of Zorithoxylum heitzii in natural forest in Cain- ZANTHOXYLUMTslHANiMPosAH. Ferner eroon. Harvesting The minimum hole diameter for Protologue Mein. ACad. Sci. (Paris) 67: 2 harvesting Is 60 cm in Garnero0n and 50 cm in (1948) Central African Republic. Family Rutaceae Handling after harvest The wood is sus- Origin and geographic distribution Zori- ceptible to blue stain and logs should be re- thoxy!urn tsihonimposo is endemic to western moved from the forest immediately after felling Madagascar, where it occurs from Nitsiranana or be treated with an anti-sapstain chemical to Morondava Logs float in water and can be transported by Uses The wood, often traded as 'fahavalon- river kazo', is used locally for house building, espe- Genetic resources Zorithoxylum heitziiis cially for window frames and doors, and also in valued for its multipurpose timber and there is shipbuilding and for boxes and crates. It is high demand locally for its bark, which is used suitable for veneer and plywood production. A in traditional medicine. Although this species bark decoction is used to treat malaria Is not considered of conservation concern at Properties The heartwood is yellowish present, the logging and debarking may make brown, sometimes with a greenish or golden it liable to genetic erosion in the near future shine, and distinctly demarcated from the grey- The pressure on the trees because of bark col- ish white sapwood. The grain is straight, tex- Iection can be locally high, as was recorded ture coarse around the Dja reserve in southern Cameroon. The wood is medium-weight, with a density of Prospects Zorithoxylum heitsit may have 520-680 kg/ina at 12% moisture content. It good prospects as a plantation timber tree, but seasons rapidly with little or no degrade. The more research Is needed on propagation tech- rates of shrinkage are moderately high, from niques. It seems to be a suitable candidate for green to oven dry 4.3-5.5% radial and 7.9-8.9% sustainable timber exploitation in natural for- tangential. Once dry, the wood is moderately est because it Is able to regenerate in logged- stable in service. At 12% moisture content, the over forest. modulus of rupture is 120-161 N/min2 The medicinal properties deserve more atten- modulus of elasticity 12,100-16,800 N/mm2 tion, particularly the analgesic activity. Sev- compression parallel to grain 46-59 N/mm2 eral of the claimed activities in traditional shear 5 N/min2, cleavage 12-13 N/min and medicine are coinparable to those of Zorithoxy- Chalais-Meudon side hardness 1.9-3.6. Ium gilletii, which has been subject to much The wood Is fairly easy to saw and work. It more pharmacologicalresearch, with promising holds nails moderately well. The gluing and results painting properties are satisfactory. The dura- Major references Bolza & Keating, 1972; hinty Is poor to moderate, the wood being sus- CIRAD Forestry Department, 2003; CTFT, ceptible to attacks by termites, Lyctus borers 1980; de Saint-Aubin, 1963; Petroff, Doat & and fungi. The heartwood is moderately resis- Tissot, 1967; Sallenave, 1955; Sallenave, 1964; tant to Impregnation with preservatives. Takahashi, 1978 Several alkaloids have been isolated from the