Gilbertiodendron dewevrei

Family

Caesalpiniaceae (Leguminosae - Caesalpinioideae)

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Chromosome number

n = 12

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Synonyms

Macrolobium dewevrei De Wild. (1904).

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Vernacular names

Limbali (Fr).

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Origin and geographic distribution

Gilbertiodendron dewevrei is distributed from Nigeria to DR Congo and northern Angola.

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Uses

The wood, traded as ‘limbali’, is very suitable for flooring, joinery, stairs, window frames, doors and decks of bridges. It can also be used for heavy construction including hydraulic works, interior trim, mine props, ship building, vehicle bodies, garden furniture, railway sleepers, toys, novelties, agricultural implements, draining boards and turnery. As the wood does not have special aesthetic qualities, it is not very suitable for cabinet work and fine joinery. The wood is considered unsuitable as firewood, but it is much sought after for the production of charcoal.
In DR Congo the inner bark is used for tying and for making bands for carrying baskets, whereas bark cylinders are sometimes used for making honey containers. The leaves are used for thatching and house walls. In north-eastern DR Congo houses were traditionally built from material of Gilbertiodendron dewevrei and clay only: poles for the house skeleton, small branches for the walls between rooms, string from the bark of young trees for tying, and leaves for covering the roof.
Although it has been recorded that it takes a long time to remove toxic compounds from the seeds, the roasted or boiled seeds are eaten in times of shortage in Central Africa, or they are boiled and ground into flour, which is made into porridge. The fermented seeds are ground, wrapped in leaves of Megaphrynium macrostachyum (Benth.) Milne-Redh., and roasted. The seeds are also eaten by the Baka pygmy people of Cameroon, after they have been boiled and the seed coat has been removed.
In traditional medicine in Congo, the powdered bark is taken for the treatment of dysentery and is sprinkled on wounds, the leaves are used against sterility and asthma, and to promote childbirth, whereas leaf ash is rubbed into scarifications on painful knees. In DR Congo sap from the stem is applied on whitlows, bark decoctions are drunk for the treatment of gastritis and blennorrhoea, sap expressed from the bark together with that of Tephrosia vogelii Hook.f. is used against otitis, and pounded scrapings of the dried bark are applied on burns.

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Production and international trade

‘Limbali’ wood was regularly exported from DR Congo into Belgium before the Second World War, but only occasionally afterwards. Today, its commercial exploitation is not important anymore, and fluctuating from year to year. Exports of ‘limbali’ logs from Cameroon were 50 m³ in 2000, 1770 m³ in 2006, 2380 m³ in 2007 and 260 m³ in 2008. Cameroon exported 140 m³ of sawn wood in 2004 and 80 m³ in 2006. Congo exported 345 m³ of finished ‘limbali’ products and 15 m³ of logs in 2004, and 30 m³ of logs and 17 m³ of sawn wood in 2006.

Properties

The heartwood is pale brown to dark reddish brown, darkening upon exposure; it is distinctly demarcated from the 5–10 cm wide, greyish or yellowish sapwood. The grain is straight or wavy, occasionally interlocked, texture medium to coarse. Gum ducts are sometimes present. Quarter-sawn surfaces are slightly mottled, with alternating pale and darker brown streaks.
The wood is medium-weight to heavy, with a density of 730–910 kg/m³ at 12% moisture content. It air dries slowly, with a tendency to split. Kiln drying should be done carefully to prevent distortion and checking. It takes about 12 days to kiln dry boards of 2.5 cm thick to 12% moisture content. The rates of shrinkage are moderate to high, from green to oven dry 3.2–6.4% radial and 7.4–11.1% tangential. Once dry, the wood is moderately stable to unstable in service.
At 12% moisture content, the modulus of rupture is 102–189 N/mm², modulus of elasticity 13,300–19,300 N/mm², compression parallel to grain 54–76 N/mm², shear 8.5–12 N/mm², cleavage 16. 5–20.5 N/mm, Janka side hardness 6670 N, Janka end hardness 6490 N and Chalais-Meudon side hardness 4.4–6.0.
Fresh wood saws relatively easily but slowly. The wood may have a significant blunting effect on saw blades, making the use of stellite-tipped saw teeth necessary. It works fairly well, but tends to blunt cutting edges. It moulds and planes easily. It holds nails and screws well, but pre-boring is recommended to prevent splitting. The wood paints, varnishes and glues well. It is not suitable for peeling. The wood is durable, being moderately resistant to resistant to fungal attack, moderately resistant to termites and resistant to marine borers. The sapwood is susceptible to Lyctus borers. The heartwood is resistant to impregnation with preservatives, the sapwood more permeable.
The wood contains 38–42.5% cellulose, 34. 5–36.5% lignin, 14.5–15.5% pentosan, 0.8–1.6% ash and 0.03–0.04% silica. The solubility is 0.6–7.1% in alcohol-benzene, 1.3–2.3% in hot water and 13.6–17.2% in a 1% NaOH solution.
Per 100 g edible portion seeds from DR Congo contained: water 9.4 g, energy 1478 kJ (353 kcal), protein 4.8 g, fat 0.6 g, carbohydrate 82.3 g, fibre 1.5 g and ash 1.5 g.

Adulterations and substitutes

Other Gilbertiodendron spp., such as Gilbertiodendron brachystegioides (Harms) J.Léonard, Gilbertiodendron ogoouense (Pellegr.) J.Léonard and Gilbertiodendron preussii (Harms) J.Léonard, are sometimes confused with Gilbertiodendron dewevrei and sold as ‘limbali’.

Description

Evergreen, large tree up to 45 m tall; bole branchless for up to 22 m, straight, cylindrical, up to 200(–300) cm in diameter, without buttresses; bark surface grey-brown to yellowish brown, rough, exfoliating in large, irregular scales, with brown lenticels, inner bark thick, fibrous, hard, red-brown; crown dense. Leaves alternate, pendulous, paripinnately compound with (2–)3(–5) pairs of leaflets; stipules ovate-lanceolate, fused, 2–8 cm × 1.5–4 cm, with 2 kidney-shaped appendages up to 2.5 cm long, more or less persistent; petiole thick, 0.5–1 cm long, rachis 2–25 cm long, narrowly grooved above, short-hairy; petiolules thick, 1–16 mm long; leaflets opposite, oblong to oblanceolate or elliptical, slightly oblique, 9–50 cm × 3–20 cm, basal leaflets usually smaller than terminal ones, base rounded to cordate, asymmetrical, apex obtuse to acuminate, leathery, usually glabrous, often with 2–4 small glands near the margin, lower surface densely papillose, pinnately veined with 14–25 pairs of lateral veins. Inflorescence a terminal or axillary, lax panicle 8–25 cm long, reddish hairy. Flowers bisexual, zygomorphic, 5-merous, fragrant; pedicel 2–4 cm long; bracteoles 2, ovate to elliptical, 1–2 cm long; sepals ovate-lanceolate to narrowly triangular, 5–8 mm long, purplish red, fused at base; petals unequal, 1 ovate, 1.5–3 cm × 2.5–3 cm, deeply 2-lobed, wine-red, other petals linear-lanceolate, 6–8 mm × 1–1.5 mm; fertile stamens 3, 1.5–2.5 cm long, rudimentary stamens 6, 0.5–2 mm long; ovary superior, 1-celled, style 2–3 cm long. Fruit an obliquely oblong to oblong-obovoid, flattened pod 15–30 cm × 6–10 cm, with distinct longitudinal ridge and numerous transversal veins, brownish, densely covered with short brown hairs, dehiscent with 2 woody valves, 4–6-seeded. Seeds orbicular to oblong or slightly triangular, flattened, 4–5 cm in diameter, shiny brown. Seedling with epigeal germination; hypocotyl 7–20 cm long, epicotyl 14–24 cm long; first 2 leaves opposite, with 2 large and 2 minute leaflets.

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Other botanical information

Gilbertiodendron comprises about 25 species and is restricted to tropical Africa, distributed from Guinea to DR Congo and Angola. It was formerly included in Macrolobium, which now consists of tropical American species only, and is closely related to Pellegriniodendron, which is even considered congeneric.
The wood of various other Central African Gilbertiodendron species is sometimes traded as ‘limbali’. Gilbertiodendron brachystegioides (Harms) J.Léonard is a fairly large tree up to 35(–40) m tall with a straight bole branchless for up to 20 m and up to 80 cm in diameter, distributed in Cameroon, Equatorial Guinea and Gabon. The wood, occasionally traded as ‘limbali’ although it releases a fetid smell on felling, is medium-weight with a density of about 710 kg/m³ at 12% moisture content.
Gilbertiodendron grandiflorum (De Wild.) J.Léonard is a small tree up to 15(–25) m tall, distributed in Nigeria, Cameroon, Gabon and DR Congo. In DR Congo the wood is valued for high-class cabinet work. It is also suitable for poles, piles, flooring, joinery, mine props, ship building, vehicle bodies, railway sleepers, agricultural implements and handles. Gilbertiodendron grandiflorum has been planted for shade. The wood has a density of 640–820 kg/m³ at 12% moisture content.
Gilbertiodendron grandistipulatum (De Wild.) J.Léonard is a medium-sized tree up to 30 m tall with a bole up to 80 cm in diameter, distributed in Gabon and DR Congo. The wood is quite similar to that of Gilbertiodendron dewevrei and has a density of about 840 kg/m³ at 12% moisture content.
Gilbertiodendron mayombense (Pellegr.) J.Léonard is a small to medium-sized tree up to 20 m tall with a bole up to 65 cm in diameter, distributed from Nigeria to DR Congo and Cabinda (Angola). The wood is considered suitable for construction, flooring, joinery, mine props, ship building, vehicle bodies, furniture, railway sleepers and agricultural implements. It has a density of about 770 kg/m³ at 12% moisture content.
Gilbertiodendron ogoouense (Pellegr.) J.Léonard is a medium-sized to fairly large tree up to 35 m tall with a cylindrical bole branchless for up to 19 m and up to 200 cm in diameter, distributed in Cameroon, Equatorial Guinea, Gabon, DR Congo and Cabinda (Angola). The wood is suitable for joinery and railway sleepers.

Anatomy

Wood-anatomical description (IAWA hardwood codes):
Growth rings: 1: growth ring boundaries distinct. 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); 25: intervessel pits small (4–7 μm); (26: intervessel pits medium (7–10 μm)); 29: vestured pits; 30: vessel-ray pits with distinct borders; similar to intervessel pits in size and shape throughout the ray cell; 43: mean tangential diameter of vessel lumina ≥ 200 μm; 46: ≤ 5 vessels per square millimetre; (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; 66: non-septate fibres present; 69: fibres thin- to thick-walled. Axial parenchyma: 79: axial parenchyma vasicentric; 80: axial parenchyma aliform; 81: axial parenchyma lozenge-aliform; 83: axial parenchyma confluent; (89: axial parenchyma in marginal or in seemingly marginal bands); 91: two cells per parenchyma strand; 92: four (3–4) cells per parenchyma strand. Rays: (96: rays exclusively uniseriate); 97: ray width 1–3 cells; 106: body ray cells procumbent with one row of upright and/or square marginal cells; 115: 4–12 rays per mm; (116: ≥ 12 rays per mm). Mineral inclusions: 136: prismatic crystals present; 142: prismatic crystals in chambered axial parenchyma cells.
(E. Uetimane, P.E. Gasson & E.A. Wheeler)

Growth and development

Germination is rapid, usually within 5–7 days after seed dispersal. Regeneration is abundant under shade, and seedlings tolerate dense shade for a long time. The roots are colonized by ectomycorrhizae and vesicular-arbuscular endomycorrhizae. The proportion of the roots that is colonized by ectomycorrhizae increases with age in saplings. The rapid colonization of the roots by ectomycorrhizae gives the seedlings an advantage over species that are not or more slowly colonized. In dense, undisturbed stands, the roots are not nodulating with Rhizobium, but some root nodules can be found in disturbed forest.
The average annual increase in stem diameter is 5.3 mm. Observations on 10 trees originating from Gabon and Congo show annual increases in stem diameter of 2.6–11.4 mm.
Young leaves are red and formed throughout the year. In Nigeria and Cameroon flowering is in January–April, and fruiting takes place in (July–)August–September(–October). Flowering does not always occur annually, and a large number of flowers is shed without forming fruits. Still, fruiting is usually abundant, with mast seed production every 2–4 years. Seed dispersal is ballistic, with the seeds being ejected after explosive dehiscence of the fruit.

Ecology

Gilbertiodendron dewevrei occurs below 1000 m altitude, in areas with an average annual rainfall of 1600–1900 mm and a dry season of about 2 months. It grows gregariously in humid depressions, alluvial valleys and along rivers, but also frequently in upland locations (plateaus, hill tops). On deep, well-draining sandy soils, it forms a deep tap root. On stony soils tap root formation is difficult, and the formation of lateral roots becomes prominent.
Gilbertiodendron dewevrei can form extensive, almost pure stands, sometimes covering more than 10,000 ha. In these monodominant forests, Gilbertiodendron dewevrei forms 75–88% of the basal area. The largest stands can be found in the northern and north-eastern part of the Congo basin, in DR Congo, where these forests are considered to be the climax vegetation. In DR Congo these stands typically consist of 3 layers: a continuous upper tree layer about 35 m high, almost uniformly consisting of Gilbertiodendron dewevrei, with in some places the crown of another large, heliophilous tree species (e.g. Anthonotha fragrans (Baker f.) Exell & Hillc., Dialium corbisieri Staner, Irvingia wombolu Vermoesen, Prioria oxyphylla (Harms) Breteler and Staudtia kamerunensis Warb.); a not very dense middle layer mainly consisting of young Gilbertiodendron dewevrei mixed with several rare semi-heliophytes (e.g. Diogoa zenkeri (Engl.) Exell & Mendonça, Garcinia punctata Oliv. and Synsepalum subcordatum De Wild.) and shrubs (e.g. Alchornea floribunda Müll.Arg. and Isolona thonneri (De Wild. & T.Durand) Engl. & Diels); and a discontinuous herbaceous layer of Marantaceae and Commelinaceae.
In certain regions the monodominant Gilbertiodendron dewevrei forest seems to gain terrain to the heterogeneous forests. Elsewhere, for instance in Cameroon, it seems to regress, being gradually invaded by species from the surrounding semi-deciduous forest, either under the influence of a decreasing groundwater table or as a result of human activities. The monodominant Gilbertiodendron dewevrei forests are very sensitive to forest clearing for shifting cultivation, because this permits the penetration of secondary and deciduous species into the forest. If forest clearing is too substantial and frequent, the monodominant forests may be transformed into semi-deciduous forest or degraded savanna after only 4–6 years of cultivation. Near the northern and southern limits of its distribution area, Gilbertiodendron dewevrei does not form stands as extensive as in the northern Congo basin, but it persists along rivers. Where Gilbertiodendron dewevrei is found in heterogeneous forest, it is never as isolated trees, but always in groups.

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Propagation and planting

Gilbertiodendron dewevrei regenerates abundantly under natural conditions. For planting, seeds should be sown immediately after collection. The 1000-seed weight is 18–26 kg. Seedlings can be planted out after 9–12 months in the nursery.

Management

In natural forest, suppression of the herbaceous layer makes the establishment of Gilbertiodendron dewevrei seedlings even more abundant. Although the species is typically shade-loving, moderate light is beneficial for the growth of seedlings, but full light has adverse effect. Excessive opening of the canopy of dense Gilbertiodendron dewevrei stands will favour the establishment of light-loving species with rapid growth, often accompanied by abundant growth of lianas, which is not desirable for the establishment of exploitable Gilbertiodendron dewevrei stands.

Diseases and pests

Fallen seeds of Gilbertiodendron dewevrei are often severely attacked by insects and are eaten by various mammals, such as antelopes, wild pigs, elephants, rodents and primates. Fungi only attack seeds already damaged by insects. Forest buffaloes and bongos eat the seedlings, and elephants sometimes dig up saplings over large areas to eat the root mat consisting of roots, fungal mycelia and decaying leaves. Primates eat the young leaves of adult trees. The young leaves of seedlings and young trees (less than 2 m tall) are most liable to damage. Adult trees do not seem to have important enemies, but they are sometimes attacked by the fungus Fomes lignosus.

Harvesting

The minimum bole diameter for felling in Cameroon (2001) and DR Congo is 60 cm, in Gabon 70 cm and in the Central African Republic 90 cm. To make headbands for carrying baskets, the Mbuti pygmy people in DR Congo strip bark strips 2 m long and 5–10 cm wide from young trees with a bole diameter of 15–20 cm, and remove the outer bark. The harvest of bark strips from young plants can lead to heart rot of trees.

Yield

In DR Congo a tree 25 m tall with a bole branchless for 10 m and a diameter of 90 cm had a wood volume of 5.5 m³. In the Uele region in DR Congo, dense stands dominated by Gilbertiodendron dewevrei had on average 419 trees with a bole diameter over 10 cm per ha, with a total volume of 370 m³/ha, of which 245 Gilbertiodendron dewevrei trees/ha with a volume of 283 m³/ha.

Handling after harvest

Gilbertiodendron dewevrei logs must be sawn soon after felling because they tend to split. Because the density of fresh wood is high, logs cannot be transported by floating.

Genetic resources and breeding

Gilbertiodendron dewevrei is not much exploited for its wood, and it is present in protected areas, sometimes in important populations. Currently there are no important threats for Gilbertiodendron dewevrei, but this may rapidly change if its wood becomes more valuable, because clear-cutting of extensive monospecific stands could, in the long run, bring about a drastic decline of populations and important genetic erosion of the species.

Breeding

No planting or breeding programmes of Gilbertiodendron dewevrei are known to exist.

Prospects

The promotion of Gilbertiodendron dewevrei wood by a number of forest exploiters having it in their concessions, among their customers as well as on Internet, could eventually raise the question of the durability of the resource. The stands rich in Gilbertiodendron dewevrei are very sensitive to invasion by other species when the canopy is opened too much. Clear-cutting these stands would be deleterious for their survival, and control of the size of the gaps created by felling should be promoted to prevent invasion of light-loving species, and to stimulate regeneration and optimal growth of Gilbertiodendron dewevrei. Ringing of the trees before felling can help to decrease the impact of falling trees. In certain more densely populated rural areas, forest clearing for agriculture can present an important danger because it brings about degradation of the monodominant Gilbertiodendron dewevrei forests, with penetration of pioneer species. Short rotations and intensive weeding of crops, as is practised in Cameroon, can lead to invasion of herbs and even to savanna formation. Therefore it would be good to monitor the evolution of Gilbertiodendron dewevrei stands to prevent these dangers from becoming realities. More research is still needed to establish appropriate silvicultural methods.

Major references

• ATIBT (Association Technique Internationale des Bois Tropicaux), 1986. Tropical timber atlas: Part 1 – Africa. ATIBT, Paris, France. 208 pp.
• Aubréville, A., 1970. Légumineuses - Césalpinioidées (Leguminosae - Caesalpinioideae). Flore du Cameroun. Volume 9. Muséum National d’Histoire Naturelle, Paris, France. 339 pp.
• Bolza, E. & Keating, W.G., 1972. African timbers: the properties, uses and characteristics of 700 species. Division of Building Research, CSIRO, Melbourne, Australia. 710 pp.
• CTFT (Centre Technique Forestier Tropical), 1960. Résultats des observations et des essais effectués au Centre Technique Forestier Tropical sur le limbali (Gilbertiodendron dewevrei). Information technique No 72, CTFT, Nogent-sur-Marne, France. 5 pp.
• Fouarge, J., Gérard, G. & Sacré, E., 1953. Bois du Congo. Institut National pour l’Etude Agronomique du Congo belge (INEAC), Brussels, Belgium. 424 pp.
• Gérard, P., 1960. Etude écologique de la forêt dense à Gilbertiodendron dewevrei dans la région de l’Uele. Série scientifique No 45, Institut National pour l’Etude Agronomique du Congo (INEAC), Brussels, Belgium. 159 pp.
• Gérard, J., Edi Kouassi, A., Daigremont, C., Détienne, P., Fouquet, D. & Vernay, M., 1998. Synthèse sur les caractéristiques technologiques des principaux bois commerciaux africains. Document Forafri 11. Cirad, Montpellier, France. 185 pp.
• Hart, T.B., 1995. Seed, seedling and sub-canopy survival in monodominant and mixed forests of the Ituri Forest, Africa. Journal of Tropical Ecology 11(3): 443–459.
• Hart, T.B., Hart, J.A. & Murphy, P.G., 1989. Monodominant and species-rich forests of the humid tropics: causes for their co-occurrence. The American Naturalist 133(5): 613–633.
• Vivien, J. & Faure, J.J., 1985. Arbres des forêts denses d’Afrique Centrale. Agence de Coopération Culturelle et Technique, Paris, France. 565 pp.

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Other references

• Ankei, T., 1990. Cookbook of the Songola: an anthropological study on the technology of food preparation among a Bantu-speaking people of the Zaïre forest. African Study Monographs, Supplement 13(1): 1–174.
• Blake, S. & Fay, J.M., 1997. Seed production by Gilbertiodendron dewevrei in the Nouabalé Ndoki National Park, Congo, and its implications for large mammals. Journal of Tropical Ecology 14: 885–891.
• Burkill, H.M., 1995. The useful plants of West Tropical Africa. 2nd Edition. Volume 3, Families J–L. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 857 pp.
• Dudek, S., Förster, B. & Klissenbauer, K., 1981. Lesser known Liberian timber species. Description of physical and mechanical properties, natural durability, treatability, workability and suggested uses. GTZ, Eschborn, Germany. 168 pp.
• Fouarge, J., Quoilin, J. & Roosen, P., 1970. Essais physiques, mécaniques et de durabilité de bois de la République Démocratique du Congo. Série technique No 76. Institut National pour l’Etude Agronomique du Congo (INEAC), Brussels, Belgium. 40 pp.
• Keay, R.W.J., 1989. Trees of Nigeria. A revised version of Nigerian trees (1960, 1964) by Keay, R.W.J., Onochie, C.F.A. & Stanfield, D.P. Clarendon Press, Oxford, United Kingdom. 476 pp.
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• Likunde, B., 1987. Contribution à l’étude floristique de la forêt à Gilbertiodendron dewevrei (De Wild.) J.Léonard de Yalisombo (Kisangani). Mémoire de licence inédit, Université de Kisangani, Faculté des Sciences, Kisangani, DR Congo. 68 pp.
• Louis, J. & Fouarge, J., 1949. Essences forestières et bois du Congo. Fascicule 6. Macrolobium dewevrei. Institut National pour l’Etude Agronomique du Congo belge (INEAC), Brussels, Belgium. 44 pp.
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• Sallenave, P., 1955. Propriétés physiques et mécaniques des bois tropicaux de l’Union française. Centre Technique Forestier Tropical, Nogent-sur-Marne, France. 129 pp.
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Afriref references

Sources of illustration

• Aubréville, A., 1968. Légumineuses - Caesalpinioidées (Leguminosae - Caesalpinioideae). Flore du Gabon. Volume 15. Muséum National d’Histoire Naturelle, Paris, France. 362 pp.
• Vivien, J. & Faure, J.J., 1985. Arbres des forêts denses d’Afrique Centrale. Agence de Coopération Culturelle et Technique, Paris, France. 565 pp.

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Author(s)

•	C. Doumenge CIRAD, Campus International de Baillarguet TA-C-105/D, F-34398 Montpellier Cédex 5, France

Editors

•	R.H.M.J. Lemmens PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
•	D. Louppe CIRAD, Département Environnements et Sociétés, Cirad es-dir, Campus international de Baillarguet, TA C 105 / D (Bât. C, Bur. 113), 34398 Montpellier Cédex 5, France
•	A.A. Oteng-Amoako Forestry Research Institute of Ghana (FORIG), University P.O. Box 63, KNUST, Kumasi, Ghana

Associate editors

•	E.A. Obeng Forestry Research Institute of Ghana (FORIG), University P.O. Box 63, KNUST, Kumasi, Ghana

Photo editor

•	G.H. Schmelzer PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands

Correct citation of this article

Doumenge, C., 2012. Gilbertiodendron dewevrei (De Wild.) J.Léonard. [Internet] Record from PROTA4U. Lemmens, R.H.M.J., Louppe, D. & Oteng-Amoako, A.A. (Editors). PROTA (Plant Resources of Tropical Africa / Ressources végétales de l’Afrique tropicale), Wageningen, Netherlands. <http://www.prota4u.org/search.asp>. Accessed .

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