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Tieghemella heckelii (A.Chev.) Roberty

Protologue
Petite Flore de l’ouest-Africain: 79 (1954).
Family
Sapotaceae
Chromosome number
2n = 24
Synonyms
Dumoria heckelii A.Chev. (1907), Mimusops heckelii (A.Chev.) Hutch. & Dalziel (1931), Baillonella heckelii (A.Chev.) Baehni (1965).
Vernacular names
Makore (En). Makoré (Fr). Makoré (Po).
Origin and geographic distribution
Tieghemella heckelii occurs in the West African forest zone, from Sierra Leone eastwards to southern Nigeria, but not in Togo or Benin.
Uses
The wood, traded as makore or cherry mahogany, is used for furniture, exterior and interior construction, flooring, doors, vehicle frames, sports goods, railway sleepers, turnery and sculptures, and makes good and decorative veneer, often used to face plywood, especially for marine uses.
The seed kernels (cotyledons, known as ‘baco’) are rich in an edible fat known as ‘dumori butter’ or ‘makore butter’, which is locally popular as a cooking or seasoning oil and often preferred to palm oil. The fat is also applied as a pomade to the body and hair, and used in soap production. The fleshy and very sticky pulp of the fruits is sometimes used as birdlime. The bark is reportedly effective for treating blennorrhoea and toothache, and in Liberia young buds are used to treat snake bites.
Production and international trade
Makore is traded on the international timber market, but production is small due to limited supply from natural stands. It is often traded together with ‘douka’ wood (from Tieghemella africana Pierre). The export of makore timber from Côte d’Ivoire declined from 70,000 m³ in 1960 to 28,000 m³ at the beginning of the 1980s and 6000 m³ at the end of the 1980s. In 1994 Ghana exported 2100 m³ of air-dried makore sawnwood for an average price of US$ 510/m³, and 3200 m³ of sliced veneer for an average price of US$ 780/m³, rotary-peeled veneer for US$ 450/m³ and jointed veneer for US$ 1735/m³. Côte d’Ivoire exported 200 m³ of veneer for US$ 1800/m³ in 1994. In 2001 Ghana exported 3500 m³ of makore veneer.
Properties
The data on wood properties of makore and douka cannot be separated in the literature, and the following description applies to both species. The wood resembles African mahogany (Khaya and Entandrophragma spp.), but the texture is finer. The heartwood is pinkish-, purplish- or reddish-brown with a silky lustre, often with a decorative figure in the form of flames or stripes; it is very distinctly demarcated from the up to 8 cm wide and pinkish white to greyish brown sapwood. The grain is straight or interlocked, texture fine and even.
The wood is medium-weight with a density of 600–800 kg/m³ at 12% moisture content. The shrinkage rates are low to moderate. In a test, shrinkage of the wood from 90% to 60% air moisture content was 1.1% radial and 1.8% tangential. Shrinkage from green to 12% moisture content was 3.0% radial and 4.5% tangential, and from green to oven dry 5.3–6.5% radial and 7.3–8.7% tangential. Usually, drying does not cause problems, although the wood dries slowly.
At 12% moisture content, the modulus of rupture is 96–165 N/mm², modulus of elasticity 9500–13,900 N/mm², compression parallel to grain 47–67 N/mm², shear 8–14 N/mm², cleavage 17–27 N/mm and Janka side hardness 4940 N.
The wood is somewhat difficult to work due to the presence of silica; blunting effects are moderately severe when sawing, and stellite-tipped sawteeth are recommended. The wood finishes well. Staining and polishing give good results. Pre-boring for nailing and screwing is recommended to avoid splitting. Gluing properties are good. The timber can be peeled satisfactorily.
The heartwood is rated as one of the most durable African timbers. It is resistant to termites and fungi. Damage by pinhole borers and powder-post beetles has been recorded occasionally. Although it is resistant to marine borers in temperate waters, the wood is not very durable in tropical (especially brackish) waters. The heartwood is very difficult to impregnate, the sapwood moderately easy.
Dust from sawn wood may cause irritation to skin and mucous membranes. It has been suggested that this is caused by the presence of saponins or the contact allergen 2,6-dimethoxy-1,4-benzoquinone. A highly haemolytic saponin has been isolated from the wood; on hydrolysis it gave d-glucose, l-rhamnose and d-xylose.
The fruit is sticky and juicy, with an unpleasant smell and bitter taste. The kernel comprises about 60% oil by weight. The oil is yellowish and semi-fluid, has no distinct flavour or taste (occasionally slightly spicy), and consists of about 51% oleic acid, 43% stearic acid, 3.5% palmitic acid and 2.5% linoleic acid.
Adulterations and substitutes
Makore is used for similar purposes as African mahogany (Khaya and Entandrophragma spp.), but it is more durable. It is sometimes even traded as African mahogany. It is very similar to the wood of Tieghemella africana from Central Africa, which is often also traded as makore or douka.
Description
Very large tree up to 55 m tall; bole up to 250 cm in diameter, sometimes much more, straight and cylindrical, often swollen in lower part, reaching up to 30 m to the first branches, angular or ridged at base, or with large, narrow buttresses up to 3 m high and heavy, spreading surface roots; bark c. 2 cm thick, surface grey to nearly black, deeply furrowed with rectangular scales, inner bark reddish, fibrous but brittle, exuding a sticky latex; crown heavy, rounded, heavy branches often abruptly spreading. Leaves arranged spirally, more or less in tufts at the ends of branches, simple; stipules absent; petiole 1.5–4 cm long, slender; blade elliptical to obovate, 6–15 cm × 2–6.5 cm, cuneate at base, emarginate, rounded, acute or acuminate at apex, with entire to slightly undulate margin, papery or thinly leathery, glabrous, lateral veins numerous. Flowers in fascicles of 1–4 (as a rule 2) in the leaf axils, bisexual, regular; pedicel 1.5–2.5 cm long; calyx with 2 whorls of 4 lobes 3–5 mm long, softly hairy on the overlapping parts; corolla with 2–2.5 mm long, fleshy tube and 8 lobes, each lobe with 1 minute median segment and 2 large, fleshy, imbricate lateral segments 3–3.5 mm long, creamy white; stamens 8, inserted on the corolla tube in front of the corolla lobes, free, filaments short, 8 short staminodes alternating with the stamens; ovary superior, conical, softly hairy, 8-celled, each cell with 1 ovule, style short. Fruit a large, ovoid-globose, smooth berry 8–12 cm long, yellow when ripe, containing 1–3 seeds in a yellowish pulp. Seeds broadly ellipsoid, slightly laterally compressed, (5–)6–7.5 cm long, testa thick, woody, smooth, shining and yellowish-brown in dorsal part, rough and bullate in ventral part (scar); endosperm absent. Seedling with epigeal germination, hypocotyl stout, 6–14 cm long, epicotyl up to 3–11 cm long, cotyledons thick, sessile, c. 5 cm × 2 cm, dark green.
Other botanical information
The genus Tieghemella 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 and thin cotyledons. Tieghemella africana resembles Tieghemella heckelii and may be conspecific. The former differs in the longer, filiform median segment of the corolla lobes, larger staminodes and slightly smaller seed scar. A taxonomic study is needed to clarify the species limits and the status of the genus, which is complicated by the fact that the name Tieghemella was first published for a genus of fungi.
Anatomy
Wood-anatomical description (IAWA hardwood codes):
Growth rings: 2: growth ring boundaries indistinct or absent. Vessels: 5: wood diffuse-porous; 7: vessels in diagonal and/or radial pattern; (10: vessels in radial multiples of 4 or more common); 13: simple perforation plates; 22: intervessel pits alternate; 23: shape of alternate pits polygonal; 25: intervessel pits small (4–7 μm); 26: intervessel pits medium (7–10 μm); (30: vessel-ray pits with distinct borders; similar to intervessel pits in size and shape throughout the ray cell); 31: vessel-ray pits with much reduced borders to apparently simple: pits rounded or angular; 32: vessel-ray pits with much reduced borders to apparently simple: pits horizontal (scalariform, gash-like) to vertical (palisade); (33: vessel-ray pits of two distinct sizes or types in the same ray cell); 42: mean tangential diameter of vessel lumina 100–200 μm; 47: 5–20 vessels per square millimetre; 56: tyloses common; (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: 86: axial parenchyma in narrow bands or lines up to three cells wide; 87: axial parenchyma reticulate; 93: eight (5–8) cells per parenchyma strand; 94: over eight cells per parenchyma strand. Rays: 97: ray width 1–3 cells; 100: rays with multiseriate portion(s) as wide as uniseriate portions; 107: body ray cells procumbent with mostly 2–4 rows of upright and/or square marginal cells; 108: body ray cells procumbent with over 4 rows of upright and/or square marginal cells; 115: 4–12 rays per mm. Mineral inclusions: 159: silica bodies present; 160: silica bodies in ray cells.
(L.N. Banak, P. Détienne & P.E. Gasson)
Growth and development
Initial growth of seedlings is fast, up to 70 cm in 4 months, but then often ceases while a strong taproot (c. 20 cm long) is formed. Growth of young makore trees is reported as slow, but strongly depends on light. Under conditions above 40% of full light, growth may be up to 1 m/year; under 10% growth is almost nil. Other reports indicate, however, that maximum growth is reached at an irradiance of 10% of unshaded values. In Ghana, young trees have been reported to be 1–2 m tall after 1–1.5 years in a nursery, and when planted under shade trees in the field 1.5–3.5 m tall after 13 years. However, in western Côte d’Ivoire, trees have been reported to be 3 m tall after 4 years and 9–11 m tall after 20 years with a stem diameter of 13–16 cm, occasionally even 28 m tall after 21 years with a diameter of 37 cm. For 35–80- year-old trees the mean annual increment is 0.4–0.8 cm.
Trees develop according to Aubréville’s model: the monopodial trunk shows rhythmic growth, with whorled branches, which also grow rhythmically but modularly, each branch plagiotropic by apposition, the modules growing indefinitely.
Trees start flowering and fruiting after about 17 years, but sometimes after 10 years. Flowers open in the early morning, the corolla being shed in the afternoon of the same day. In Liberia, flowering is from February to May; ripe fruits can be found between October and December. In Côte d’Ivoire, flowering occurs from January to June, and ripe fruits can be found from August to March. A large tree produces approximately 3000–4000 fruits at a time. The fruits are eaten by elephants, which are probably the main seed dispersers; bush-pigs are also reported to feed on the fruits.
Ecology
Tieghemella heckelii is often an emergent tree of the high forest. It is found in moist evergreen and semi-deciduous forests. Regeneration in these forests is reportedly reasonable, at least in Ghana; there is a strong preference for undisturbed forest. Young trees are very tolerant of shade, but can survive in full sun. Heavy soils are preferred.
Propagation and planting
Makore is easy to propagate. The seeds are heavy, 30–50 seeds/kg. Viability decreases rapidly. They should be planted within 2 weeks. In the nursery, seeds are usually planted at a spacing of 40 cm × 40 cm under light shade. Seedbeds should be watered well. Germination is over 90% in optimal conditions, and without pre-treatment. It begins after about 4 weeks, most seeds having germinated after 10 weeks, but it may continue for 16 weeks. Rodents may cause considerable damage to seeds and seedlings (cotyledons). Saplings of about 2.5 years old and 1–1.5 m tall are planted out into the field usually at the beginning of the rainy season, but in western Côte d’Ivoire 0.5 year-old seedlings are used.
Propagation by air layering and cuttings is successful. In a test with air layering, 39% of branches rooted after 8 weeks. The average rate of rooting of branch cuttings after 6 months was 73%. In neither case did the application of indole-butyric acid improve rooting success.
Management
The density of Tieghemella heckelii in the forest is generally low. Reports for Côte d’Ivoire vary from 1 exploitable tree per 23 ha to 1.4 trees greater than 10 cm diameter per ha. The average of 9 inventories in Côte d’Ivoire was 4.1 m³ of timber per 100 ha for trees exceeding 70 cm diameter. The average of 16 inventories in Liberia was 33 m³ of timber per 100 ha for trees exceeding 40 cm diameter. Experiments in Côte d’Ivoire showed that thinning of the forest leads to better recruitment and growth.
Makore has been planted on a very small scale (almost 6 ha), mainly for seed oil production, in western Côte d’Ivoire (near Taï National Park) with an average density of 357 stems/ha. It is often planted at a density of about 120 trees/ha, in association with coffee, cocoa, rubber or rice, and at field margins and in secondary forest following clearance of undergrowth.
Diseases and pests
No information on diseases and pests is available, although Tieghemella heckelii is a host of the semi-parasitic tree Okoubaka aubrevillei Pellegr. & Normand.
Harvesting
Logs tend to split badly during felling. Logs of larger trees may be hollow.
Yield
Fat yield is 20–30 kg/tree per harvest. One kg of seed kernels yields about 200 g of fat when the traditional method of extraction is used.
Handling after harvest
Care should be taken when sawing makore wood because nasal and respiratory irritation may occur. To extract the oil, the seeds are split, and the kernels dried in the sun, roasted, pounded to a paste and boiled in water. The fat or oil is skimmed off from the water.
Genetic resources
Tieghemella heckelii is reported to be rather rare in Liberia. The density in Côte d’Ivoire is low, and in many areas it is considered rare. In Ghana, it is fairly common, but under pressure as a result of forest exploitation, and deserves protection. The seeds are dispersed by elephants, which are becoming rare in West Africa and therefore limiting natural regeneration.
Tieghemella heckelii is included in the IUCN Red list of threatened species because it suffers both from habitat destruction and selective logging practices.
Prospects
The prospects for planting programmes in West Africa using Tieghemella heckelii are good. Seed and vegetative propagation is easy. The species can be planted in open sites, grows fairly rapidly, and has a fairly open crown structure which permits good light penetration, making it suitable for use in agroforestry programmes. The wood, and fat from the seeds, are of excellent quality. However, planting for fat production may be uneconomic as it takes a rather long time for trees to fruit after planting, and fat yields are limited. Planting is needed to take pressure off wild populations.
Major references
• Ayensu, E.S. & Bentum, A., 1974. Commercial timbers of West Africa. Smithsonian Contributions to Botany 14. Smithsonian Institution Press, Washington D.C., United States. 69 pp.
• Bonnéhin, L., 2000. Domestication paysanne des arbres fruitiers forestiers. Cas de Coula edulis Baill., Olacaceae, et de Tieghemella heckelii Pierre ex A. Chev., Sapotaceae, autour du Parc National de Taï, Côte d’Ivoire. Tropenbos-Côte d’Ivoire Série 1. 138 pp.
• Burkill, H.M., 2000. The useful plants of West Tropical Africa. 2nd Edition. Volume 5, Families S–Z, Addenda. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 686 pp.
• CIRAD Forestry Department, 2003. Makore. [Internet] Tropix 5.0. http://tropix.cirad.fr/ afr/makore.pdf. Accessed July 2004.
• CTFT (Centre Technique Forestier Tropical), 1976. Makoré - Douka. Bois et Forêts des Tropiques 168: 31–46.
• Heine, H., 1963. Sapotaceae. In: Hepper, F.N. (Editor). Flora of West Tropical Africa. Volume 2. 2nd Edition. Crown Agents for Oversea Governments and Administrations, London, United Kingdom. pp. 16–30.
• Richter, H.G. & Dallwitz, M.J., 2000. Commercial timbers: descriptions, illustrations, identification, and information retrieval. [Internet]. Version 18th October 2002. http://delta-intkey.com/wood/index.htm. Accessed July 2004.
• Siepel, A., Poorter, L. & Hawthorne, W.D., 2004. Ecological profiles of large timber species. In: Poorter, L., Bongers, F., Kouamé, F.N. & Hawthorne, W.D. (Editors). Biodiversity of West African forests. An ecological atlas of woody plant species. CABI Publishing, CAB International, Wallingford, United Kingdom. pp. 391–445.
• Voorhoeve, A.G., 1979. Liberian high forest trees. A systematic botanical study of the 75 most important or frequent high forest trees, with reference to numerous related species. Agricultural Research Reports 652, 2nd Impression. Centre for Agricultural Publishing and Documentation, Wageningen, Netherlands. 416 pp.
Other references
• Agyeman, V.K., Swaine, M.D. & Thompson, J., 1999. Responses of tropical forest tree seedlings to irradiance and the derivation of a light response index. Journal of Ecology 87: 815–827.
• Bokdam, J., 1977. Seedling morphology of some African Sapotaceae and its taxonomical significance. Mededelingen Landbouwhogeschool Wageningen 77–20. Wageningen, Netherlands. 84 pp.
• Durrieu de Madron, L., Favrichon, V., Dupuy, B., Bar-Hen, A. & Maître, H.-F., 1998. Croissance et productivité en forêt dense humide: bilan des expérimentations dans le dispositif d’Irobo, Côte d’Ivoire (1978–1990). Document Forafri 2. Cirad, Montpellier, France. 69 pp.
• Gosse, B., Gnabre, J., Bates, R.B., Dicus, C.W., Nakkiew, P. & Huang, R.C.C., 2002. Antiviral saponins from Tieghemella heckelii. Journal of Natural Products 65(12): 1942–1944.
• Hawthorne, W.D., 1995. Ecological profiles of Ghanaian forest trees. Tropical Forestry Papers 29. Oxford Forestry Institute, Department of Plant Sciences, University of Oxford, United Kingdom. 345 pp.
• InsideWood, undated. [Internet] http://insidewood.lib.ncsu.edu/search/. Accessed May 2007.
• Takahashi, A., 1978. Compilation of data on the mechanical properties of foreign woods (part 3) Africa. Shimane University, Matsue, Japan. 248 pp.
• van Rompaey, R.S.A.R., 1993. Forest gradients in West Africa. A spatial gradient analysis. PhD thesis, Wageningen Agricultural University, Wageningen, Netherlands. 142 pp.
• Wiselius, S.I., 1994. Hout-vademecum. 7th edition. Stichting Centrum Hout, Almere, Netherlands. Kluwer Techniek. 380 pp.
Sources of illustration
• Voorhoeve, A.G., 1979. Liberian high forest trees. A systematic botanical study of the 75 most important or frequent high forest trees, with reference to numerous related species. Agricultural Research Reports 652, 2nd Impression. Centre for Agricultural Publishing and Documentation, Wageningen, Netherlands. 416 pp.
Author(s)
L. Bonnéhin
Projet autonome pour la conservation du Parc national de Taï (PACPNT), 01 B.P. 693, San Pédro, Côte d’Ivoire
R.H.M.J. Lemmens
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands


Editors
D. Louppe
CIRAD, Département Environnements et Sociétés, Cirad es-dir, Campus international de Baillarguet, TA C-DIR / B (Bât. C, Bur. 113), 34398 Montpellier Cedex 5, France
A.A. Oteng-Amoako
Forestry Research Institute of Ghana (FORIG), University P.O. Box 63, KNUST, Kumasi, Ghana
M. Brink
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
General editors
R.H.M.J. Lemmens
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
L.P.A. Oyen
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
J.R. Cobbinah
Forestry Research Institute of Ghana (FORIG), University P.O. Box 63, KNUST, Kumasi, Ghana
Photo editor
E. Boer
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands

Correct citation of this article:
Bonnéhin, L. & Lemmens, R.H.M.J., 2005. Tieghemella heckelii (A.Chev.) Roberty. In: Louppe, D., Oteng-Amoako, A.A. & Brink, M. (Editors). Prota 7(1): Timbers/Bois d’œuvre 1. [CD-Rom]. PROTA, Wageningen, Netherlands.

















































Distribution Map wild


1, flowering twig; 2, flower; 3, fruit; 4, seed; 5, seed kernel
Redrawn and adapted by W. Wessel-Brand



tree habit, trees protected by local people


base of bole


14-year-old tree


leaves and fruit


fruits and seeds


fruit, seeds and germinating seed


rooted branches 8 weeks after air layering


planting in rice field


wood (radial face)


wood (tangential face)


wood in transverse section


wood in tangential section


wood in radial section