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Terminalia tetrandra (Danguy) Capuron

Protologue
Bull. Mus. natl. Hist. nat., sér. 3, bot. 11: 97 (1973).
Family
Combretaceae
Synonyms
Terminaliopsis tetrandrus Danguy (1923).
Origin and geographic distribution
Terminalia tetrandra is endemic to eastern Madagascar, occurring from Sambava to the south.
Uses
The wood, known as ‘tafanala’ is used for joinery, furniture and tool handles. It is suitable for light construction, flooring, interior trim, ship building, vehicle bodies, musical instruments, boxes, crates, toys, novelties, vats, turnery, veneer, plywood, hardboard and particle board.
The bark is used in traditional medicine; it is taken to treat excessive salivation, and is applied as a wash to boils and ulcers. It is also used in the production of alcoholic drinks with a bitter taste. It is sometimes planted as an ornamental tree.
Properties
The heartwood is pinkish pale brown, with uneven yellowish markings. It is clearly demarcated from the yellowish, 5–7 cm wide sapwood. The grain is slightly wavy, sometimes interlocked, texture moderately coarse.
The wood is medium-weight, with a density of 560–690(–750) kg/m³ at 12% moisture content. It air dries well, with little degrade. The rates of shrinkage are moderate, from green to oven dry 3.1–4.8% radial and 6.9–9.3% tangential. Board of 2.5 cm thick take 2–3 months to air dry to 30% moisture content.
At 12% moisture content the modulus of rupture is 118–148 N/mm², modulus of elasticity 8820–12,300 N/mm², compression parallel to grain 50–68 N/mm², shear 6–8 N/mm², cleavage 14.5–17.5 N/mm and Chalais-Meudon side hardness 3.1–5.1.
The wood is easy to saw and work with both hand and machine tools, with moderate blunting effect on cutting edges. The wood finishes well, but the use of a filler is necessary to obtain an acceptable polish. It holds nails and screws well, and glues moderately well. The wood is not durable and susceptible to attacks by fungi, termites and Lyctus borers. It is difficult to impregnate with preservatives. Wood splinters may cause inflammation of the skin.
Adulterations and substitutes
The wood of Terminalia tetrandra resembles that of Terminalia superba Engl. & Diels from mainland Africa, which is sometimes planted in Madagascar.
Description
Deciduous, medium-sized to fairly large tree up to 35 m tall; bole branchless for up to 12 m, usually straight, up to 100(–120) cm in diameter; inner bark yellow; crown storied with branches in whorls, spreading; branchlets slender, very young twigs with minute hairs. Leaves arranged spirally, clustered near ends of branchlets, simple; stipules absent; petiole up to 2.5 cm long, slender; blade obovate to elliptical-obovate, up to 10 cm × 4 cm, long-cuneate at base, rounded to short-acuminate at apex, margins finely toothed, leathery, glabrous, pinnately veined with 7–12 pairs of indistinct lateral veins. Inflorescence an axillary spike 4–10 cm long, slender; peduncle 2–5 cm long. Flowers bisexual or male, regular, 4–5-merous, up to 5 mm long, yellowish; receptacle spindle-shaped; sepals triangular; petals absent; stamens 4–5, free, long-exserted; disk annular, densely hairy; ovary inferior, 1-celled, style thick, narrowly conical. Fruit an ellipsoid drupe c. 2 cm × 1 cm, slightly 4–5-angled, smooth and glabrous, greenish, crowned by the remains of the sepals and disk, indehiscent, 1-seeded.
Other botanical information
Terminalia is a pantropical genus of about 200 species. In tropical mainland Africa about 30 species occur naturally, in Madagascar about 35. Terminalia tetrandra differs from other Terminalia spp. in having only 4–5 stamens in a single whorl (versus 8–10 in 2 whorls), and it has been placed in a separate genus (Terminaliopsis) for this reason. However, as this is not accompanied by other characters, it is considered insufficient for genus status.
Anatomy
Growth rings: (1: growth ring boundaries distinct); (2: growth ring boundaries indistinct or absent). Vessels: 5: wood diffuse-porous; (12: solitary vessel outline angular); 13: simple perforation plates; 22: intervessel pits alternate; 23: shape of alternate pits polygonal; 26: intervessel pits medium (7–10 μm); 27: intervessel pits large ( 10 μm); 29: vestured pits; 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. Tracheids and fibres: 61: fibres with simple to minutely bordered pits; 66: non-septate fibres present; 69: fibres thin- to thick-walled. Axial parenchyma: 80: axial parenchyma aliform; 81: axial parenchyma lozenge-aliform; 83: axial parenchyma confluent; (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; 94: over eight cells per parenchyma strand. Rays: 97: ray width 1–3 cells; 104: all ray cells procumbent; 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.
(E. Uetimane, H. Beeckman, P. Détienne & P. Gasson)
Growth and development
The tree shows sympodial growth, with new lateral shoots developing from buds in the leaf axils. These shoots grow rapidly and characteristically exist of a long, slender, leafless basal part and a short apical part with clustered leaves. This growth model results in a crown with tiers of branches, but old trees may lose the storied structure of the crown. Terminalia tetrandra mainly flowers from December to January. In the basal part of the inflorescence usually bisexual flowers develop, in the apical part male flowers.
Ecology
Terminalia tetrandra occurs in more humid evergreen forest, from sea-level up to 1100 m altitude. It seems to be rather uncommon.
Handling after harvest
Freshly harvested logs should be removed from the forest rapidly, or de-barked and treated with fungicides and insecticides, to avoid attacks by fungi and borers.
Genetic resources
Although Terminalia tetrandra is widely distributed in eastern Madagascar, it is rather uncommon and limited to a threatened habitat of moist, dense, evergreen forest. It may easily become threatened by genetic erosion because it is selectively logged for its timber.
Prospects
Little is known about Terminalia tetrandra, particularly concerning its ecological preferences, growth rates and regeneration. Research is warranted because of the existence of successful timber plantations of other Terminalia spp., e.g. Terminalia superba. Terminalia tetrandra may be a serious candidate for the establishment of timber plantations in Madagascar.
Major references
• 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.
• Capuron, R., 1966. Etudes sur les essences forestières de Madagascar : tafanala (Terminalia tetrandra - Combretaceae). Centre Technique Forestier Tropical, Section de Madagascar, Tananarive, Madagascar. 5 pp.
• Capuron, R., 1973. Contribution à l’étude de la Flore forestière de Madagascar. Notes sur le genre Terminalia L. Bulletin du Muséum Nationale d’Histoire Naturelle, sér. 3, botanique 11: 89–179.
• Guéneau, P., 1971. Bois de Madagascar. Possibilités d’emploi. Centre Technique Forestier Tropical, Antananarivo, Madagascar. 75 pp.
• Guéneau, P., Bedel, J. & Thiel, J., 1970–1975. Bois et essences malgaches. Centre Technique Forestier Tropical, Nogent-sur-Marne, France. 150 pp.
• Perrier de la Bâthie, H., 1954. Combrétacées (Combretaceae). Flore de Madagascar et des Comores (plantes vasculaires), familles 147–151. Firmin-Didot et cie., Paris, France. 84 pp.
• Sallenave, P., 1964. Propriétés physiques et mécaniques des bois tropicaux. Premier supplément. Centre Technique Forestier Tropical, Nogent-sur-Marne, France. 79 pp.
• Sallenave, P., 1971. Propriétés physiques et mecaniques des bois tropicaux. Deuxième supplément. Centre Technique Forestier Tropical, Nogent-sur-Marne, France. 128 pp.
• Takahashi, A., 1978. Compilation of data on the mechanical properties of foreign woods (part 3) Africa. Shimane University, Matsue, Japan, 248 pp.
Other references
• Anonymous, 1962. Tafanala (Terminaliopsis tetrandrus). Informations techniques no 160, Centre Technique Forestier Tropical, Nogent- sur-Marne, France. 1 pp.
• Boiteau, P., Boiteau, M. & Allorge-Boiteau, L., 1999. Dictionnaire des noms malgaches de végétaux. 4 Volumes + Index des noms scientifiques avec leurs équivalents malgaches. Editions Alzieu, Grenoble, France.
• InsideWood, undated. [Internet] http://insidewood.lib.ncsu.edu/search/. Accessed May 2007.
• Neuwinger, H.D., 2000. African traditional medicine: a dictionary of plant use and applications. Medpharm Scientific, Stuttgart, Germany. 589 pp.
• Novy, J.W., 1997. Medicinal plants of the eastern region of Madagascar. Journal of Ethnopharmacology 55: 119–126.
• Schatz, G.E., 2001. Generic tree flora of Madagascar. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 477 pp.
Sources of illustration
• Schatz, G.E., 2001. Generic tree flora of Madagascar. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 477 pp.
Author(s)
R.H.M.J. Lemmens
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands


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-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

Correct citation of this article:
Lemmens, R.H.M.J., 2009. Terminalia tetrandra (Danguy) Capuron. In: Lemmens, R.H.M.J., Louppe, D. & Oteng-Amoako, A.A. (Editors). Prota 7(2): Timbers/Bois d’œuvre 2. [CD-Rom]. PROTA, Wageningen, Netherlands.
Distribution Map wild


1, flowering branch; 2, bisexual flower; 3, male flower; 4, fruits.
Redrawn and adapted by Iskak Syamsudin