Prota 7(1): Timbers/Bois d’œuvre 1
Mém. Mus. natl. Hist. nat., Paris 19: 250 (1830).
2n = 50
Dry-zone mahogany, Senegal mahogany, Gambia mahogany (En). Acajou du Sénégal, acajou caïlcédrat (Fr). Acaju do Senegal, bisselon, mogno de Africa (Po).
Origin and geographic distribution
Khaya senegalensis occurs from Mauritania and Senegal east to northern Uganda. It is commonly planted within its natural area of distribution, mainly as ornamental and roadside tree, and also outside this area, e.g. in Cape Verde, Tanzania, Malawi, Madagascar, Réunion, Egypt, South Africa, India, Indonesia, Vietnam, Australia and tropical America. In drier zones in Sri Lanka it has become a priority species for timber plantation establishment since mid 1990s, with more than 500 ha of plantation established by 2004 and 200 ha/year of new plantations planned for the future.
The wood is valued for carpentry, joinery, furniture, cabinet work, ship building and decorative veneer. It is suitable for construction, flooring, interior trim, vehicle bodies, toys, novelties, railway sleepers, turnery and pulpwood. Traditionally, the wood is used for dug-out canoes, household implements such as mortars and spoons, and drums. It is also used as fuelwood and for charcoal production.
The bitter-tasting bark is highly valued in traditional medicine. Bark decoctions or macerations are widely taken against fever caused by malaria, and against stomach complaints, diarrhoea, dysentery and anaemia, as anodyne in cases of rheumatism and headache, and as tonic, emmenagogue and anthelmintic. They are also used as purgative, antidote and abortifacient, and to treat syphilis, leprosy, chickenpox and angina. The bark is applied externally as disinfectant in cases of inflammations and to treat skin diseases, rash, scabies, wounds, ulcers, boils, haemorrhoids, swellings and toothache. The bark is commonly used in veterinary medicine, as anthelmintic, tonic and appetizer, and to treat trypanosomiasis, liver flukes, diarrhoea and ulcers. In Uganda it is used as fish poison. In Cameroon the bark is in demand as an additive in local beer brewing. Leaves are also used in traditional medicine, to treat skin complaints, wounds, jaundice, oedema, headache and depression, and as purgative. Roots are applied against jaundice, stomach-ache, oedema and amenorrhoea. The roots and/or bark are an ingredient of complex arrow poisons of which Strophanthus roots or seeds are the main ingredients. Flowers are used in medicines against stomach complaints and syphilis. Seed oil is rubbed in to treat rheumatism and influenza, and it is taken to treat syphilis. Young twigs and roots are used as chewing sticks and toothbrushes.
In Ghana the bark has been used for dyeing cloth brownish. The foliage is a common source of fodder, but it has a low fodder quality and is mainly used towards the end of the dry season when better-quality forage is not available, or in mixtures with better fodders. The seed oil is used in cosmetics and for cooking. The wood ash is added to stored grain to prevent insect attack. Khaya senegalensis is commonly planted as a roadside tree and ornamental shade tree, and sometimes for soil stabilization. It has been planted successfully in Burkina Faso in a taungya system with groundnut as intercrop. In many regions it is considered a magic tree used in rituals.
Production and international trade
Logs of Khaya senegalensis have been exported from West Africa already since the first half of the 19th century, e.g. from Gambia. Khaya senegalensis has been heavily exploited for its timber since then. Nowadays, the wood is mainly used locally, and statistics on production and trade are not available. In several countries in the African savanna area Khaya senegalensis wood is very important, e.g. in Burkina Faso and Mali, where it may contribute up to 80% of all logs entering local sawmills. Probably the wood of Khaya senegalensis is occasionally mixed with the wood of other Khaya spp. and traded on the international timber market. The bark is in high demand for medicinal purposes and can be found on many local markets. Seeds are harvested from natural stands and traded worldwide, e.g. 400–600 kg of seed is distributed per year from the Centre National de Semences Forestières (CNSF) in Burkina Faso, of which more than 80% is exported.
The heartwood is pinkish brown, darkening to reddish brown with a purplish tinge upon exposure. It is usually distinctly demarcated from the paler, up to 8 cm wide sapwood, at least in dried wood. The grain is usually interlocked, sometimes straight, texture moderately coarse.
The wood is moderately heavy, with a density of (620–)710–810(–900) kg/m³ at 12% moisture content. It generally air dries rather slowly, but with little degrade; the presence of tension wood may cause splitting and warping. The rates of shrinkage are moderate, from green to oven dry 4.0–5.9% radial and 4.3–7.2% tangential. Once dry, the wood is fairly stable in service.
At 12% moisture content, the modulus of rupture is 82–122 N/mm², modulus of elasticity (7200–)9800–11,650 N/mm², compression parallel to grain 45–54(–72) N/mm², cleavage 15–28 N/mm and Chalais-Meudon side hardness 3.5–5.9(–7.2).
The wood is fairly easy to saw and work, with moderate blunting effect on cutting edges. However, surfaces tend to become woolly and cutting edges should be kept sharp. A reduced cutting angle is recommended when machining quarter-cut boards. The wood holds nails and screws well. It has good gluing properties. It takes polishes and paints well, but the use of a filler is necessary. The peeling properties are poor because of the higher density in comparison with Khaya anthotheca and Khaya ivorensis and the interlocked grain, but slicing gives decorative veneer. The heartwood is moderately durable to durable, being resistant to termites but moderately susceptible to fungi. The sapwood is susceptible to Lyctus. The heartwood is strongly resistant to impregnation, the sapwood moderately resistant. The gross energy value of the wood is about 19,990 kJ/kg.
Bark extracts showed in-vitro antiviral activity. They also showed in-vitro antibacterial properties against strains of Enterococcus faecalis and Streptococcus sp. They exhibited significant leishmanicidal activity. The bark showed anticonvulsant effect in mice, and weak antimalarial activity in mice inoculated with Plasmodium berghei. Bark extracts exhibited pronounced antiplasmodial activity against Plasmodium falciparum strains, with IC50 values less than 5 μg/ml. Some limonoids isolated from the bark showed in-vitro antimalarial activity against strains of Plasmodium falciparum, e.g. fissinolide, which also has molluscicidal properties. The anthelmintic effects of the bark have been confirmed in in-vitro tests as well as in in-vivo tests in sheep. In tests with rats, the bark showed anti-inflammatory activity after local application. Aqueous extracts of the bark and leaves exhibited strong antisickling activity. The main active constituent was identified as a rearranged limonoid, of which the activity was much higher than that of pentoxifylline used as standard in managing sickle cell disease. In addition, it did not alter significantly the corpuscular indices. Bark extracts displayed antiproliferative, anti-inflammatory and pro-apoptotic effects on human colorectal cancer cell lines. The limonoid 3α,7α-dideacetylkhivorin isolated from the bark showed significant growth-inhibitory activity against several cancer cell lines. The extracts showed larvicidal effect against the mosquito Culex annulirostris, comparable to azadirachtin, a well-known insecticide of plant origin. Some limonoids isolated from the bark showed significant antifeedant and growth-inhibitory activities against the cotton leafworm Spodoptera littoralis; khayanolide B showed strongest activity. Polyphenols from the bark have high antioxidant activity. Leaves contain in general about 5 g of digestible protein and 620 kJ net energy per 100 g dry matter, signifying poor fodder quality. Leaves (moisture content 67.2%) from the Sudanian zone of West Africa were found to contain 8.2% crude protein and 3.7% digestible protein, on a dry matter basis. Leaf extracts are highly toxic to rice weevil (Sitophilus oryzae) and may have potential as protectant in stored grain.
The seeds have an oil content of up to 67% and are rich in oleic acid. However, seed oil contents of only 17–27% have also been recorded. Seeds from Senegal contained 58.5% oil, with as principal fatty acids: oleic acid (70.3%), linoleic acid (10.8%), palmitic acid (8.3%) and stearic acid (8.3%).
Adulterations and substitutes
The wood of Khaya senegalensis resembles true mahogany (from Swietenia spp.) more closely than the wood of Khaya anthotheca and Khaya ivorensis do, but it is heavier and harder. It most closely resembles the wood of Khaya grandifoliola C.DC. The wood of makore (Tieghemella) is similar, but more durable.
More or less evergreen, monoecious, medium-sized tree up to 30(–35) m tall; bole branchless for up to 10(–16) m but often much shorter and crooked, up to 100(–250) cm in diameter, buttresses short or absent; bark surface grey to dark grey or greyish brown, initially smooth but becoming scaly with thin, rounded scales, inner bark dark pink to reddish, exuding a reddish gum; crown rounded, dense; twigs glabrous. Leaves arranged spirally but clustered near ends of branches, paripinnately compound with (2–)3–5(–6) pairs of leaflets; stipules absent; petiole and rachis together up to 25 cm long; petiolules 3–4 mm long; leaflets opposite or nearly so, elliptical to oblong, 5–12 cm × 2.5–5 cm, cuneate and slightly asymmetrical at base, obtuse or very shortly acuminate at apex, margins entire, thinly leathery, glabrous, pinnately veined with 8–10 pairs of lateral veins. Inflorescence an axillary or seemingly terminal panicle up to 20 cm long. Flowers unisexual, male and female flowers very similar in appearance, regular, usually 4-merous, whitish, sweet-scented; pedicel 1–2 mm long; calyx lobed almost to the base, with rounded lobes c. 1.5 mm long; petals free, elliptical, c. 4 mm × 2 mm, somewhat hooded; stamens fused into an urn-shaped tube c. 5 mm long, with usually 8 included anthers near apex, alternating with rounded lobes; disk cushion-shaped; ovary superior, globose to conical, 1–2 mm in diameter, 4-celled, style up to 1 mm long, stigma disk-shaped; male flowers with rudimentary ovary, female flowers with smaller, non-dehiscing anthers. Fruit an erect, nearly globose, woody capsule 4–6 cm in diameter, pale grey to greyish brown, dehiscent by 4 valves, many-seeded. Seeds disk-shaped or quadrangular, strongly flattened, c. 2 cm × 2.5 cm, narrowly winged all around the margin, brown. Seedling with hypogeal germination, cotyledons remaining enclosed in the seed coat; epicotyl 5–6 cm long; first 2 leaves opposite, simple.
Other botanical information
Khaya comprises 4 species in mainland Africa and 1 or 2 endemic to the Comoros and Madagascar. It belongs to subfamily Swietenoideae and seems most closely related to Carapa and Swietenia. Khaya species strongly resemble each other in flowers and fruits, and differences are most prominent in their leaflets. Khaya senegalensis is close to Khaya anthotheca (Welw.) C.DC. and Khaya grandifoliola C.DC., but is usually smaller in all parts.
Wood-anatomical description (IAWA hardwood codes):
Growth rings: (1: growth ring boundaries distinct); (2: growth ring boundaries indistinct or absent). Vessels: 5: wood diffuse-porous; 13: simple perforation plates; 22: intervessel pits alternate; (23: shape of alternate pits polygonal); 24: intervessel pits minute (≤ 4 μm); 30: vessel-ray pits with distinct borders; similar to intervessel pits in size and shape throughout the ray cell; 42: mean tangential diameter of vessel lumina 100–200 μm; 47: 5–20 vessels per square millimetre; 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; 69: fibres thin- to thick-walled. Axial parenchyma: 78: axial parenchyma scanty paratracheal; 79: axial parenchyma vasicentric; (89: axial parenchyma in marginal or in seemingly marginal bands); 92: four (3–4) cells per parenchyma strand; 93: eight (5–8) cells per parenchyma strand. Rays: 98: larger rays commonly 4- to 10-seriate; 106: body ray cells procumbent with one row of upright and/or square marginal cells; 107: body ray cells procumbent with mostly 2–4 rows of upright and/or square marginal cells; 115: 4–12 rays per mm. Secretory elements and cambial variants: 131: intercellular canals of traumatic origin. Mineral inclusions: 136: prismatic crystals present; 137: prismatic crystals in upright and/or square ray cells.
(N.P. Mollel, P. Détienne & E.A. Wheeler)
Growth and development
Seedlings develop a taproot in deeper soils. They may reach a height of 10 cm after 3 months, with roots about 25 cm long. Under natural conditions, initial growth is slow, seedlings being 12–25 cm tall after 2 years. Early growth may be much faster, but Khaya senegalensis is often attacked by Hypsipyla shoot borers and it is browsed by cattle and other herbivores, resulting in slow growth and poor form of the stem. Under favourable and unshaded conditions, with the topsoil loosened, seedlings reached 0.5–1.2 m in height after 2 years, in Nigeria even 2 m with a stem diameter of 4.7 cm after 2 years. In very favourable conditions in Senegal even higher growth rates have been recorded. In Côte d’Ivoire Khaya senegalensis trees planted in the open in the semi-deciduous forest zone reached an average height of 9 m and an average bole diameter of 16.5 cm after 10 years. In the savanna of northern Côte d’Ivoire trees reached a mean height of 5.5 m at 7.5 years after planting, and sprouts reached 4.8 m after 4.5 years. In southern Burkina Faso, on deep soils on river banks, Khaya senegalensis has a diameter up to 50 cm after 25 years. In Benin the average bole diameters in plantations of about 48 years old varied between 30.5 cm and 52 cm. In planted trees in Australia, exceptional annual growth rates of 3 m in height and 4 cm in bole diameter have been recorded.
Trees usually gradually lose their leaves in the dry season, the fallen leaves often immediately being replaced by new ones. Flowering occurs at the end of the dry season or beginning of the rainy season. The flowers are pollinated by insects such as bees and moths. Fruits mature 3–5 months after flowering. Trees can start producing seeds after 20–25 years. Dispersal of the seeds is by wind, but most seeds fall close to the parent tree, normally up to 100 m.
Khaya senegalensis occurs in savanna woodland, often in moist localities and along watercourses, in areas with 650–1300(–1800) mm annual rainfall and a dry season of 4–7 months. It occurs up to 1500(–1800) m altitude. In riparian forest it can sometimes be found together with Khaya grandifoliola. It prefers deep and well-drained alluvial soils and termite mounds, but can also be found on shallow, rocky soils, where it usually remains much smaller. It tolerates flooding in the rainy season.
Propagation and planting
Natural regeneration can be abundant in savanna areas on good soils without too much competition from herbs, and when protected from fire. In the first years, seedlings tolerate light shade.
The 1000-seed weight is 140–330 g. The seeds are often attacked by insects while they are still on the tree, and undamaged seeds should therefore be selected before sowing. Soaking for 24 hours in water reportedly improves germination, but is not necessary. The seeds can best be sown in seed beds in the nursery or in pots. Fresh healthy seeds have a high germination rate, 90–100%, and they may retain their viability for 6(–8) months. However, when they are exposed to high relative humidity, they may lose their viability within 3 months. When stored at 0–10°C and a seed moisture content of 5%, seeds retain their high germination rate for at least 4 years. However, other tests showed that storage at 4°C resulted in loss of viability between 12 and 18 months, whereas storage at –20°C, 15°C and 20°C all showed good results. It is recommended to add ash during storage to reduce insect attacks.
Upon sowing seeds should be covered with only a thin layer of soil, or left partially uncovered. Germination takes 10–18 days. It has been recommended to provide light shade to young seedlings until they are 1–2 months old. In Mali and Côte d’Ivoire seedlings are planted out when they are 3–4 months old and 25–30 cm tall. Seedlings can also be left in the nursery for about one year until they are 0.5–1 m tall, after which the root system is pruned to about 30 cm long and most leaves stripped off before planting out into the field. Stumps with 2–3 cm of stem and 25–30 cm of root can also be planted out. In Senegal 50% of the stumps planted out into the field survived after 5 years, but for good survival rates regular watering after planting is needed. To reduce damage by grazing, seedlings may be planted out when over 1.5 m tall. Normal spacing is 4–5 m × 4–5 m. Wildlings are sometimes collected for planting. Trees also reproduce by root suckers. Grafting and layering is possible, but propagation using cuttings is much more difficult.
In northern Côte d’Ivoire pure plantations were unsatisfactory because of numerous attacks of by Hypsipyla shoot borers from as early as the 2nd year after plantation onwards. Planting of small plots with 5–9 Khaya senegalensis transplants 1 m apart from each other within a plantation of teak trees planted at the same time significantly reduced borer attacks and allowed good initial growth of both species. Planting Khaya senegalensis at wide spacing under light cover in a thinned natural forest or plantation also reduced attacks.
In general, Khaya senegalensis occurs scattered under natural conditions, often as single individuals. Enrichment planting in woodland has been applied successfully in Vietnam. Plantations in northern Togo established in 1918 showed 70 years later an average tree height of only 12 m and an average bole diameter of 32 cm; possibly they failed due to poor silvicultural management. In Benin the first plantations were established in 1935, but also here they usually failed, not only because of poor management, but probably also because of illegal logging of the largest individuals.
The application of a complete fertilizer at a rate of 200 g/tree at the time of planting is recommended. In young plantations weeding is necessary, as young trees are susceptible to suppression by weeds. Young trees are also susceptible to fire, but older trees are quite fire resistant. In Côte d’Ivoire Khaya senegalensis has been planted under the shade of 2-year-old Leucaena leucocephala (Lam.) de Wit, which suppresses weeds and fixes nitrogen into the soil. Regular thinning of the shade trees in the first years is needed for good growth of the Khaya senegalensis trees. The latter reached an average height of 4.9 m after 5 years, which was more than trees planted nearby in the open, with an average height of 4.0 m. In tropical Africa Khaya senegalensis has been planted successfully in mixed plantations with Azadirachta indica A.Juss., Senna siamea (Lam.) Irwin & Barneby and Dalbergia sissoo Roxb. ex DC., in Benin also in a mixture with teak (Tectona grandis L.f.).
Realistic rotation cycles under natural conditions are probably in the range of 80–100 years, but in plantations a rotation of 40–60 years is feasible.
Khaya senegalensis trees planted along streets often grow faster than in those in forest plantations, but pruning is necessary to obtain a nice bole.
Diseases and pests
In plantations Khaya senegalensis suffers seriously from Hypsipyla robusta shoot borers that kill the main stem of young trees, causing excessive branching and contributing to mortality. Silvicultural techniques such as overhead shading of saplings, mixed planting and removal of lateral shoots can reduce damage by shoot borers. Products based on methidathion have proved effective in plantations of up to 2 years old, but the costs are very high. In Burkina Faso roadside trees have been attacked by leaf-eating caterpillars, e.g. of Bourgognea microcera. Seeds are commonly attacked by seed-boring beetles and eaten by small rodents, whereas young plants can be heavily browsed by cattle, antelopes and other herbivores.
The logs are quite difficult to fell using traditional equipment because of the dense and fairly hard wood. Firewood is normally collected from fallen branches, as cross-cutting and splitting of larger dimensions of wood is difficult. Bark is collected whenever needed, and in many regions many larger trees show signs of debarking. In some regions the crowns are heavily affected by harvesting the branches for fodder.
In experimental plantations in Burkina Faso the annual production has been estimated at 3.7 m³/ha. In dry forests in northern Côte d’Ivoire a tree 51 cm in diameter yields on average 1.4 m³ of timber, and a tree 67 cm in diameter 2.6 m³.
Khaya senegalensis is included in the IUCN Red list as a vulnerable species because of habitat loss and degradation, and selective felling for its timber. Like other Khaya spp., populations have been depleted in many regions through centuries of commercial exploitation. The large-scale harvesting of the bark for medicinal purposes and of branches for fodder constitutes another threat to Khaya senegalensis populations. It has been reported for some regions in Benin that Fulani people harvest the full crown of more than 70% of the trees, and that additionally the bark is commonly collected as malaria medicine. The harvesters prefer larger trees, and heavily exploited populations showed significantly lower densities of seedlings and saplings than populations under less pressure.
In-situ conservation stands for seed production have been identified and managed in partnership with local people by the Centre National de Semences Forestières (CNSF) in Burkina Faso. The most comprehensive provenance trial reported so far has been established in the early 1970s near Darwin, Australia, with provenances from 9 African countries.
The actual overexploitation of Khaya senegalensis for timber, fodder and medicine is a serious threat for many of its populations. Therefore, sustainable methods of harvesting should be established and implemented as soon as possible. However, much research is still needed to realize this, whereas the environmental conditions and traditional land use are complications that should be considered. The fair growth rate under appropriate conditions makes extensive establishment of plantations an option, but Hypsipyla attack is a serious drawback. The combined effects of selection of provenances with genetic resistance and appropriate silvicultural practices could have a substantial positive impact on the damage caused by Hypsipyla robusta stem borers. Research priority should be given to range-wide selection of genotypes which are resistant to stem-borer attack, are fast growing and have acceptable wood quality. The establishment of appropriate methods of vegetative propagation including tissue culture is urgently needed.
The bark demonstrated several interesting pharmacological activities, such as antimalarial, anti-inflammatory and anticancer effects. This deserves more research attention for possible development into new drugs. The insecticidal and anthelmintic activities of the bark are also noteworthy.
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Correct citation of this article:
Nikiema, A. & Pasternak, D., 2008. Khaya senegalensis (Desr.) A.Juss. In: Louppe, D., Oteng-Amoako, A.A. & Brink, M. (Editors). Prota 7(1): Timbers/Bois d’œuvre 1. [CD-Rom]. PROTA, Wageningen, Netherlands.
1, tree habit; 2, flowering twig; 3, fruits; 4, seed.
Redrawn and adapted by Iskak Syamsudin
tree habit on termite hill
bark and slash
13-year-old plantation, Côte d'Ivoire
pruned tree, Burkina Faso
leaves, fruits and seeds
seeds in fruit
market under Khaya senegalensis, Burkina Faso
harvesting of bark
drying of planks
wood in transverse section
wood in tangential section
wood in radial section