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Greenwayodendron suaveolens (Engl. & Diels) Verdc.

Protologue
Adansonia, sér. 2, 9: 90 (1969).
Family
Annonaceae
Synonyms
Polyalthia suaveolens Engl. & Diels (1901).
Vernacular names
Molinda (En). Moambe noir (Fr). Muamba preta (Po).
Origin and geographic distribution
Greenwayodendron suaveolens is widespread from southern Nigeria east to western Uganda and northern Tanzania, and south to southern DR Congo and Cabinda (Angola).
Uses
The wood is used for house construction, joinery, mine props, furniture, stakes for yam cultivation, rafters and shafts of spears. It is suitable for flooring, interior trim, railway sleepers, toys, novelties, agricultural implements, vats, draining boards, food containers, turnery, veneer and plywood. The wood burns brightly and is used for illumination.
Various plant parts are used in traditional medicine. Bark decoctions are taken to treat stomach-ache and other pains, gonorrhoea and infertility, as diuretic, purgative and aphrodisiac, and to facilitate childbirth. Bark ash is rubbed into scarifications on the forehead to treat psychosis, and bark pulp is applied externally against rheumatism, headache, epilepsy and toothache. In Cameroon bark is applied to scarifications to treat malaria, and also in Gabon the bark is used for the treatment of malaria. Root decoctions are taken to treat liver complaints and headache, and root sap is administered as anthelmintic and aphrodisiac, and to treat oedema and swollen glands. Leaf decoctions or macerations serve to treat hepatitis and pains, and are applied externally to treat rheumatism. In DR Congo pounded bark is used in a mixture with other plants to make arrow poison. The fruit is edible.
Properties
The heartwood is yellow to brown when dry and usually not distinctly demarcated from the sapwood, which is yellowish white when freshly cut, but darkening upon exposure. The grain is usually straight, texture variable. Quarter-sawn surfaces show streaky markings. The wood is lustrous.
The wood is medium-weight, with a density of 750–790 kg/m³ at 12% moisture content. It air dries well, but with a slight risk of checking and end splitting. At 12% moisture content, the modulus of rupture is 151–170 N/mm², modulus of elasticity 17,450–19,800 N/mm², compression parallel to grain 63–71 N/mm², cleavage 14–17.5 N/mm and Chalais-Meudon side hardness 4.4–4.7.
The wood is easy to saw and work, both with machine and hand tools. It can be planed to a smooth and lustrous surface. The nailing properties are satisfactory, with good holding power. The wood glues well, and the steam bending properties are good. The wood is only moderately durable, being liable to termite, Lyctus and marine borer attacks. It is easy to impregnate with preservatives.
Several alkaloids have been isolated from the bark, including some indolosesquiterpenes and aporphines. Oliverine showed filaricidal activity against Onchocerca volvulus, and polycarpol against Onchocerca gutturosa. Methanolic bark and leaf extracts showed in-vitro cytotoxic activity in human monocytes, as well as antileishmanial and antifungal activities.
The major compounds in the leaf oil are α-humulene (34%) and β-caryophyllene (33%), and in the fruit oil myrcene (34%).
Description
Deciduous, medium-sized to fairly large tree up to 35(–45) m tall; bole branchless for up to 25 m, straight, cylindrical, up to 70(–90) cm in diameter, sometimes grooved at base; bark surface smooth, grey to blackish, often with hoop-marks, inner bark fibrous, yellow to orange or pale brown, becoming brownish or blackish upon exposure, with strong resinous smell; crown dense, conical, with horizontal branches; young twigs yellowish hairy, soon becoming glabrous. Leaves alternate, simple and entire; stipules absent; petiole 2–7 mm long; blade elliptical to oblong-elliptical, 4–12(–28) cm × 1.5–5.5(–10) cm, rounded to cuneate at base, acuminate at apex, papery to slightly leathery, nearly glabrous, pinnately veined with 5–13 pairs of lateral veins. Inflorescence an up to 8(–12)-flowered fascicle, often opposite the leaves, short-hairy. Flowers bisexual or male, regular; pedicel 3–9 mm long; sepals 3, fused at base, broadly ovate to nearly round, 2–3.5 mm long, short-hairy outside; petals in 2 whorls of 3, free, linear-oblong, 1–3 cm long, finely hairy, yellow to greenish white; stamens numerous in male flowers and up to 12 in bisexual flowers, linear, 1–4 mm long; carpels 12–20, linear-oblong, c. 2.5 mm long. Fruit consisting of up to 10(–13) indehiscent, ellipsoid to globose follicles 1–2 cm long, stipe 5–8 mm long, purplish red to bluish purple when ripe, 1–2(–3)-seeded. Seeds depressed globose, c. 1 cm in diameter, warty-wrinkled, with a groove, endosperm ruminate.
Other botanical information
Greenwayodendron comprises 2 species and is restricted to tropical Africa. It has been separated from Polyalthia, which is a genus of about 120 species, most of them occurring in tropical Asia and Australia, about 15 in Madagascar and 3 in East Africa. Although wood anatomical characteristics are quite similar, molecular studies showed that Greenwayodendron is probably not closely related to Polyalthia, supporting a status as separate genus.
Subsp. usambaricum Verdc. has been distinguished as a distinct subspecies endemic to the Usambara Mountains in Tanzania. Subsp. suaveolens var. gabonica (Pellegr. ex Le Thomas) Verdc. is endemic to Gabon and has been distinguished because of its larger leaves, which are slightly short-hairy below, and larger flowers. Chloroplast DNA studies showed distinct genetic divergence, indicating that the two sympatric varieties are most probably reproductively isolated and might represent true biological species.
Greenwayodendron oliveri (Engl.) Verdc. (synonym: Polyalthia oliveri Engl.) is a small tree up to 15 m tall occurring in rainforest from Sierra Leone to Ghana. The boles are used in house building. The yellowish brown wood is rather heavy, hard and strong. Bark decoctions and infusions are taken to treat blackwater fever and stomach complaints, and the bark is also used as vermifuge.
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)); 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; 46: 5 vessels per square millimetre; 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; 70: fibres very thick-walled. Axial parenchyma: 78: axial parenchyma scanty paratracheal; 86: axial parenchyma in narrow bands or lines up to three cells wide; 88: axial parenchyma scalariform; 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; 99: larger rays commonly > 10-seriate; 102: ray height > 1 mm; (103: rays of two distinct sizes); 104: all ray cells procumbent; (106: body ray cells procumbent with one row of upright and/or square marginal cells); 114: 4 rays per mm; 115: 4–12 rays per mm. Secretory elements and cambial variants: (124: oil and/or mucilage cells associated with ray parenchyma).
(C. Essien, P.E. Gasson & E.A. Wheeler)
Growth and development
Greenwayodendron suaveolens is classified as shade tolerant. In Gabon the fruits ripen in November–March. The fruit pulp is eaten by monkeys and chimpanzees, which may disperse the seeds. Elephants and hornbills have also been recorded as seed dispersers.
Ecology
Greenwayodendron suaveolens occurs in humid evergreen and semi-deciduous forest, often as an understorey tree. In Uganda it occurs up to 1100 m altitude. It is considered a typical component of mature forest.
Management
In Cameroon the average number of boles of over 15 cm in diameter has been recorded as 2.6 per ha, with an average wood volume of 1.8 m³/ha.
Harvesting
In Cameroon and Gabon the minimum bole diameter for harvesting is 60 cm, in the Central African Republic and DR Congo 70 cm.
Yield
A bole of 70 cm in diameter and 10 m long yielded 3.8 m³ of wood.
Genetic resources
Greenwayodendron suaveolens is widespread and is locally a common understorey tree. Therefore, it does not seem to be threatened at present, but with the ongoing decline in primary rainforest, it might become under pressure in the near future.
Prospects
Little information is available on this species, but in view of its often limited bole size and probable slow growth rates as understorey tree, it does not seem to have good prospects as commercially valuable timber tree. Interesting pharmacological activities have been demonstrated and these deserve more research attention.
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.
• Burkill, H.M., 1985. The useful plants of West Tropical Africa. 2nd Edition. Volume 1, Families A–D. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 960 pp.
• Dauby, G., Duminil, J., Heuertz, M. & Hardy, O.J., 2010. Chloroplast DNA polymorphism and phylogeography of a Central African tree species widespread in mature rainforests: Greenwayodendron suaveolens (Annonaceae). Tropical Plant Biology 3: 4–13.
• le Thomas, A., 1969. Annonacées. Flore du Gabon. Volume 16. Muséum National d’Histoire Naturelle, Paris, France. 372 pp.
• Tailfer, Y., 1989. La forêt dense d’Afrique centrale. Identification pratique des principaux arbres. Tome 2. CTA, Wageningen, Pays Bas. pp. 465–1271.
• Takahashi, A., 1978. Compilation of data on the mechanical properties of foreign woods (part 3) Africa. Shimane University, Matsue, Japan. 248 pp.
• Vivien, J. & Faure, J.J., 1985. Arbres des forêts denses d’Afrique Centrale. Agence de Coopération Culturelle et Technique, Paris, France. 565 pp.
• Wilks, C. & Issembé, Y., 2000. Les arbres de la Guinée Equatoriale: Guide pratique d’identification: région continentale. Projet CUREF, Bata, Guinée Equatoriale. 546 pp.
• Williams, R.B., Hu, J.-F., Olson, K.M., Norman, V.L., Goering, M.G., O’Neil-Johnson, M., Eldridge, G.R. & Starks, C.M., 2010. Antibiotic indole sesquiterpene alkaloid from Greenwayodendron suaveolens with a new natural product framework. Journal of natural products 73(5): 1008–1011.
Other references
• Betti, J.L., 2001. Vulnérabilité des plantes utilisées comme antipaludiques dans l'arrondissement de Mintom au sud de la réserve de biosphère du Dja (Cameroun). Systematics and Geography of Plants 71: 661-678.
• Boutique, R., 1951. Annonaceae. In: Robyns, W., Staner, P., Demaret, F., Germain, R., Gilbert, G., Hauman, L., Homès, M., Jurion, F., Lebrun, J., Vanden Abeele, M. & Boutique, R. (Editors). Flore du Congo belge et du Ruanda-Urundi. Spermatophytes. Volume 2. Institut National pour l’Étude Agronomique du Congo belge, Brussels, Belgium. pp. 256–389.
• Cooper, G.P. & Record, S.J., 1931. The evergreen forests of Liberia. School of Forestry, Yale University, Bulletin 31, New Haven, United States. 153 pp.
• Cravo, L., Perineau, F., Delmas, M. & Bessière, J.M., 1991. Chemical composition of the fruit and leaf oil of Polyalthia suaveolens Engl. et Diels. Journal of Essential Oil Research 3(6): 459–461.
• Hawthorne, W. & Jongkind, C., 2006. Woody plants of western African forests: a guide to the forest trees, shrubs and lianes from Senegal to Ghana. Kew Publishing, Royal Botanic Gardens, Kew, United Kingdom. 1023 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.
• Lamidi, M., DiGiorgio, C., Delmas, F., Favel, A., Mve-Mba, C.E., Rondi, M.L., Ollivier, E., Nze-Ekekang, L. & Balansard, G., 2005. In vitro cytotoxic, antileishmanial and antifungal activities of ethnopharmacologically selected Gabonese plants. Journal of Ethnopharmacology 102: 185-190.
• Mols, J.B., Gravendeel, B., Chatrou, L.W., Pirie, M.D., Bygrave, P.C., Chase, M.W. & Kessler, P.J.A., 2004. Identifying clades in Asian Annonaceae: monophyletic genera in the polyphyletic Miliuseae. American Journal of Botany 91(4): 590–600.
• Nkeoua, G. & Boundzanga, G.C., 1999. Données sur les produits forestières non ligneux en République du Congo. FAO, Brazzaville, Congo. 125 pp.
• Neuwinger, H.D., 2000. African traditional medicine: a dictionary of plant use and applications. Medpharm Scientific, Stuttgart, Germany. 589 pp.
• Normand, D. & Paquis, J., 1976. Manuel d’identification des bois commerciaux. Tome 2. Afrique guinéo-congolaise. Centre Technique Forestier Tropical, Nogent-sur-Marne, France. 335 pp.
• Nyasse, B., Ngantchou, I., Nono, J.-J. & Schneider, B., 2006. Antifilarial activity in vitro of polycarpol and 3-O-acetyl aleuritolic acid from Cameroonian medicinal plants against Onchocerca gutturosa. Natural Products Research 20(4): 391–397.
• Raponda-Walker, A. & Sillans, R., 1961. Les plantes utiles du Gabon. Paul Lechevalier, Paris, France. 614 pp.
• Terashima, H. & Ichikawa, M., 2003. A comparative ethnobotany of the Mbuti and Efe hunter-gatherers in the Ituri forest, Democratic Republic of Congo. African Study Monographs 24(1–2): 1–168.
• Verdcourt, B., 1971. Annonaceae. In: Milne-Redhead, E. & Polhill, R.M. (Editors). Flora of Tropical East Africa. Crown Agents for Oversea Governments and Administrations, London, United Kingdom. 131 pp.
• White, L. & Abernethy, K., 1997. A guide to the vegetation of the Lopé Reserve, Gabon. 2nd edition. Wildlife Conservation Society, New York, United States. 224 pp.
Sources of illustration
• le Thomas, A., 1969. Annonacées. Flore du Gabon. Volume 16. Muséum National d’Histoire Naturelle, Paris, France. 372 pp.
• Wilks, C. & Issembé, Y., 2000. Les arbres de la Guinée Equatoriale: Guide pratique d’identification: région continentale. Projet CUREF, Bata, Guinée Equatoriale. 546 pp.
Author(s)
R.B. Jiofack Tafokou
Ecologic Museum of Cameroon, P.O. Box 8038, Yaoundé, Cameroon


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
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:
Jiofack Tafokou, R.B., 2011. Greenwayodendron suaveolens (Engl. & Diels) Verdc. 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, base of bole; 2, flowering branch; 3, fruiting branch; 4, seeds.
Redrawn and adapted by Iskak Syamsudin



Greenwayodendron suaveolens


Greenwayodendron suaveolens


Greenwayodendron suaveolens


Greenwayodendron suaveolens