Lophira alata

Family

Ochnaceae

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Chromosome number

2n = 28

Synonyms

Lophira procera A.Chev. (1935).

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Vernacular names

Azobé, ekki, red ironwood (En). Azobé (Fr). Azobe (Po).

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Origin and geographic distribution

The area of natural distribution of Lophira alata extends from Guinea Bissau eastward to the Central African Republic, and southward to Gabon and DR Congo. It is most common in coastal areas, less so inland; in DR Congo it occurs from the western borders to the central basin of the river Congo.

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Uses

The wood of Lophira alata, well-known in trade as ‘azobé’, has become one of the most widely used tropical hardwoods, as it is available in larger dimensions than other woods with similar technical specifications. Because of its hardness and resistance properties, it is well appreciated for heavy construction work for extreme conditions, including hydraulic works, marine construction, bridges, decks, ship building, industrial floors, staircases, wooden frame houses, railway sleepers, mine props, retaining structures, elevator guides in mines, vehicle bodies and posts for a variety of purposes from vineyards to oyster culture. It is suitable for joinery, interior trim, sporting goods, toys, novelties, agricultural implements, turnery and hardboard. A notable application is for rails for metro carriages in Paris running on pneumatic tyres.
Azobé has a high energy value. It is a good fuelwood and yields an excellent charcoal. The flowers attract honey bees.
In traditional medicine in Gabon, the bark is applied topically to treat kidney problems. In Cameroon, Gabon and Congo, bark decoctions are taken or applied as enema against menstrual problems, beginning hernia and stomach problems. They are also taken against kidney pain and toothache. In Congo powdered bark mixed with palm oil and mineral salt is administered to treat heart problems, costal pain and coughing up blood. In Nigeria a similar mixture is used against cough, fever and jaundice. The bark is credited with analgesic and sedative properties and is used to treat convulsions, epilepsy, eye problems and yaws. In the Central African Republic the bark is applied against snake bites, in Cameroon for wound healing. In Liberia and Congo the leaves are used in the treatment of leprosy, and in Liberia the seeds are sometimes used similarly. The leaves are also used as a wash for women during childbirth, as a lotion against respiratory diseases and dysentery, whereas they are also added to preparations which are administered for the treatment of yellow fever and sleeplessness. Twigs are used as toothbrush.
The seed contains about 40% oil, which is sometimes used for cooking, e.g. by the Baka Pygmy people in Cameroon, and also as ointment and to make soap. In traditional medicine, it is probably used for similar purposes as ‘meni oil’ extracted from the seeds of Lophira lanceolata Tiegh. ex Keay.

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Production and international trade

Lophira alata is one of the most exploited forest trees of tropical Africa. The earliest export record of the timber dates from the 16^th century and documents its sale from Ghana to the United Kingdom for use as keels in ship building. Already before 1914, azobé wood has been used in Cameroon in the construction of quays and pontoons, and regular export to Europe started soon after World War I.
In 2006 azobé ranked 4^th on the list of exploited timbers in Cameroon with 5.1% of the total harvested timber volume, representing about 17,500 m³. It has retained its prominence in recent years with export volumes of sawn wood reaching 48,800 m³, 44,800 m³ and 49,600 m³ in 2006, 2007 and 2008, respectively. Export volumes of logs during this period were very small. Also in Gabon and Equatorial Guinea, azobé is an important export timber, but well behind okoumé (Aucoumea klaineana Pierre). In Gabon it holds second place for locally transformed timber, with 37,700 m³ entering sawmills in 2007 (against 931,500 m³ for okoumé). From Gabon it is exported as logs in similar quantities (37,500 m³ in 2008), but holding only 7^th place among log exports. During the first 4 months of 2010, ITTO has reported price increments of several tropical woods, including azobé, due to strong demand from China. In 2010 Liberia exported logs of azobé at US$ 253 per m³.

Properties

Commercial logs range in diameter from 60 cm to 120 cm, occasionally reaching 150 cm. The heartwood is chestnut brown to chocolate brown or dark violet-brown, slightly darkening on exposure, and rather poorly demarcated from the paler, about 3(–5) cm thick sapwood. Between the sapwood and the heartwood there is a 5–10 cm thick layer of transitional wood which is intermediate in colour. The grain is usually interlocked, texture variable to coarse.
The wood is very heavy, with a density of 1010–1150 kg/m³ at 12% moisture content, and very hard. Drying should be done slowly because of a high risk of deformation, surface checking and splitting. It is generally recommended to air dry azobé in pieces of large dimensions. The rates of shrinkage are high, from green to oven dry (5.6–)7.1–9.2% radial and (8.3–)10.7–13.2% tangential. Once dry, the wood is very unstable in service.
The wood has excellent mechanical properties, being both flexible and resistant to bending and very resistant to shocks and abrasion. At 12% moisture content, the modulus of rupture is (159–)201–287 N/mm², modulus of elasticity 13,200–22,000(–23,500) N/mm², compression parallel to grain (75–)85–109 N/mm², shear (10–)14–20 N/mm², cleavage 20–33 N/mm, Janka side hardness 9990–17,540 N, Janka end hardness 12,300–21,200 N and Chalais-Meudon side hardness 7.6–18.8.
Azobé is very difficult to work with hand tools because of its hardness and high density. It is normally processed by sawing, and is not suitable for peeling or slicing for veneer. With appropriate heavy-duty machine tools it saws well. The blunting effect on saw teeth and cutting edges is relatively small as azobé does not contain silica. Azobé planes without difficulty and sands very easily, but the presence of interlocked grain makes it difficult to obtain a nicely polished surface. Pre-boring is needed for nailing and screwing, but holding properties of the wood are good. Nailing is possible with special tools such as a ramset gun, and high-quality screws are required. Azobé glues well with synthetic resin glues, but gluing needs care because of the strong shrinkage on drying, and gluing on an industrial scale is not recommended. The wood takes paint and varnish well, provided it is dry. Oil-based paints chip easily. Bending is difficult.
The heartwood is very durable, but sapwood and the transitional wood are only moderately durable. The heartwood is very resistant to termite attack, but attacks by Macrotermes spp. have been reported from Africa. Resistance of dry wood to attack by Lyctus and other insects is good, although attacks to sapwood have been reported. Resistance to wood-attacking fungi is variable, but generally good. Azobé shows satisfactory resistance against marine borers in cold and temperate waters, although less than some other tropical hardwoods. In tropical waters the resistance is only moderate. Azobé is rather resistant to impregnation with preservatives. When preservation is needed, wood should be scarified immediately after sawing and just before treatment under pressure.
Azobé has a strong resistance to fire, being almost uninflammable. It is of no value for the paper industry, being difficult to reduce to shavings and having a low yield of a dark pulp. Azobé is resistant to acids, e.g. against sulphuric and hydrochloric acids at a concentration of less than 25% at ambient temperatures. It is less resistant to nitric acid. Its hardness is not affected by acids. It is also tolerant of caustic alkaline solutions (<5%).
Several bi-flavonoids have been extracted from the bark, including lophirones B, C, D, E, L and M, bongosine, mbamichalcone and lophirochalcone. Luteolin and lithospermoside have also been isolated. Extracts of the bark have shown bactericidal effects against gram-positive bacteria including Micrococcus luteus and Staphylococcus aureus. An extract of the leaves has shown activity against the vector of schistosomiasis.
The seed oil of Lophira alata has not been studied as extensively as the oil of Lophira lanceolata, but is probably fairly similar and suitable for similar purposes.

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Adulterations and substitutes

Lophira lanceolata Van Tiegh. ex Keay is closely related to Lophira alata, has similar bactericidal effects and contains similar compounds (lophirones), but its seed oil is more widely used than that of Lophira alata. It occurs in the savanna areas north of the area of distribution of Lophira alata, and in areas where they occur side by side it is possible that the two species are utilized interchangeably. The wood of Lophira lanceolata, which is pinkish with a red heart, is available in much smaller dimensions, but may have similar uses, especially for mine props, railway sleepers and mortars, as fuelwood and for charcoal production.

Description

Deciduous, large to very large tree up to 60 m tall; bole branchless for up to 30 m, cylindrical and usually very straight but sometimes slightly wavy, up to 160(–180) cm in diameter, base sometimes slightly thickened or with slightly swollen main roots; superficial, spreading roots sometimes visible for several meters; bark surface scaly, rusty brown to orange-brown, peeling in numerous, long and narrow vertical strips, inner bark up to 2 cm thick, brown-red, with a characteristic, thin, sulphur-yellow layer on the outside between the living and dead bark layers; crown hemispherical, with large, upright branches; twigs glabrous. Leaves arranged spirally, clustered at the ends of branches, simple and entire; stipules triangular, c. 6 mm long, caducous; petiole 0.5–2.5 cm long; blade oblong-obovate, 8–24(–35) cm × 4–8(–10) cm, base cuneate, apex rounded or notched, leathery, glabrous, midrib prominent, pinnately veined with many, closely arranged, fine lateral veins. Inflorescence a lax, terminal panicle, glabrous. Flowers bisexual, nearly regular, 5-merous, strongly scented; pedicel up to 2.5 cm long, jointed just below apex; sepals free, ovate, slightly unequal, c. 1 cm long; petals free, broadly obovate to nearly round, up to 1.5 cm long, slightly 2-lobed at apex, white; stamens numerous, free, c. 1 cm long, yellow-orange; ovary superior, narrowly conical, 1-celled, style arms 2, short, spreading. Fruit a conical nut 2.5–3 cm × 1–1.5 cm, tapering at apex, somewhat woody, indehiscent, 1-seeded, with 2 unequal, narrowly oblong, pink to reddish, finely veined wings at base, the larger one up to 12 cm × 3 cm, the smaller one up to 6 cm × 1 cm. Seed narrowly ovoid. Seedling with hypogeal germination; epicotyl 8–14 cm long; cotyledons remaining in seed coat, c. 1.5 cm × 0.5 cm; the first 2 leaves opposite, sessile, narrowly elliptical, c. 10 cm × 3 cm, the 3^rd leaf (and occasionally 4^th) unfolding at the same time as the first 2, forming a pseudowhorl, subsequent leaves alternate.

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Other botanical information

Lophira is restricted to tropical Africa and comprises 2 species with an ecologically distinct distribution: Lophira alata, a large tree characteristic of the Guineo-Congolian evergreen rainforest, and Lophira lanceolata, a small tree restricted to Sudano-Guinean savanna, also at higher elevations up to 1200(–1600) m altitude. In the forest-savanna transition zone, the two species are very similar in their vegetative characteristics, but Lophira lanceolata usually has comparatively narrower leaves with a longer petiole and slightly larger fruits.

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Anatomy

Wood-anatomical description (IAWA hardwood codes):
Growth rings: 2: growth ring boundaries indistinct or absent. Vessels: 5: wood diffuse-porous; 13: simple perforation plates; 22: intervessel pits alternate; 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; 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: (60: vascular/vasicentric tracheids present); 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: 85: axial parenchyma bands more than three cells wide; 94: over eight cells per parenchyma strand. Rays: 97: ray width 1–3 cells; 104: all ray cells procumbent; 115: 4–12 rays per mm. Mineral inclusions: 136: prismatic crystals present; 141: prismatic crystals in non-chambered axial parenchyma cells; 142: prismatic crystals in chambered axial parenchyma cells.
(E. Uetimane, P. Baas & H. Beeckman)

Growth and development

Lophira alata is a heliophile pioneer species. It regenerates abundantly in open localities, especially when the soil has been disturbed, e.g. along forest roads or river banks, in forest gaps, abandoned fields, or edges of savanna protected from fire. It can also germinate in the undergrowth, but full sunlight is needed for subsequent growth of the seedling. Young plants can persist fairly long in the undergrowth without developing; their growth becomes very rapid once they are exposed to stronger light. The response is stronger when the light increases from 2% to 30% of full sunlight, then weaker between 30% and 60%, suggesting adaptation to partial shade. In plantations, young plants grow on average 80–130 cm/year in height, but growth can reach and even exceed 1.5 m per year for the most vigorous plants. In a locality in Ghana, where exploitation for timber and poles was largely abandoned, Lophira alata reached a height of 6–8 m after 9 years, while the trees were in good health in spite of being shaded by 15–18 m tall umbrella trees.
The average diameter growth rates vary strongly in natural forest. In Côte d’Ivoire growth rates of 0.65–1.5 cm per year have been reported for trees with a bole diameter of more than 10 cm; the maximum growth rate observed was 2.25 cm per year for a tree that grew from 28.6 cm to 78.5 cm in diameter in 22 years. In plantations diameter growth rates of 0.85–1.15 cm per year have been measured for trees of about 10 years old, exceeding 1.5 cm per year in the most vigorous ones. While in natural forest growth can be fast for young trees in full sunlight, growth slows down immediately when competition with other species occurs. When mature trees reach and dominate the forest canopy, they regain good growth rates. It has been estimated that trees with a bole diameter of 70 cm are at least 90 years old.
Lophira alata loses its leaves for a period of a few days to 2–3 weeks very soon after the end of the rainy season, in October–November in West Africa, in December in coastal Cameroon but slightly earlier in inland areas of Cameroon, and in December–February in Gabon. New leaves may start unfolding immediately after leaf-fall or even before leaf-fall is complete. They are reddish and may appear at different times in different parts of a single crown. Leaf-fall and the appearance of new leaves may be synchronous in all trees in a region, but can also extend over 2 months. Flowering coincides with the appearance of new leaves. Fruits develop quickly and ripen after 6–8 weeks, generally in January–April, but ripe fruits can be found on the tree also later, sometimes till May–June. Not all trees bear fruit annually, and it has been reported from Guinea that mast years occur only every 3–4 years.
Leaves of juvenile trees are much larger than those in adult trees, up to 100 cm × 10 cm, are generally lanceolate and nearly sessile, and often bear galls. In the Central African Republic, trees with a bole diameter of 15 cm have been found flowering, but only trees with a bole diameter of more than 50 cm flower annually. However, the number of fruits under a mother tree becomes markedly larger when the bole diameter has become larger than 110 cm, or when the tree avails of a large exposed crown. This indicates that sexual maturity and reproductive efficiency are reached late.
In Cameroon it has been observed that roots are colonized by arbuscular endomycorrhyzae; the degree of infestation can be a factor favouring growth.

Ecology

Lophira alata is characteristic of dense humid evergreen forest at low altitudes, but it is occasionally found up to 1000 m altitude, e.g. in Cameroon. It is abundant in sedimentary river basins along the Atlantic coast, generally on dry land, but sometimes also, but more thinly spread, in periodically flooded forest behind mangroves or along the edges of marshes, rarely inside marshes. It is often found in secondary forest and regrowth along forest roads and in clearings, and is also common in forest islands in savanna and in open Marantaceae forest in Gabon and Congo. Further inland, Lophira alata occurs more scattered, and is mainly found in valleys and on the banks of greater rivers. It can spread rather far into semi-deciduous forest in river valleys and on hill slopes. Its intolerance of drought excludes it from dry climates, except in low-lying sites where sufficient water is available. In West Africa, the abundance of Lophira alata is closely linked with rainfall, reaching an optimum at 2600 mm per year. It is adapted to various types of soils, including sands, sandy clays, and gravelly and ferralitic soils, but seems to prefer sandy clay soils with a fairly shallow water table.
Young trees are very sensitive to competition, even in forest regrowth, where other species are often more efficient, such as large herbs, shrubs, lianas and trees such as Musanga cecropioides R.Br. and Pycnanthus angolensis (Welw.) Warb.
The regeneration of Lophira alata seems to require light and the near complete absence of competition during the seedling and very young plant stages. Trees are fairly resistant to fire as long as the damage is limited to one side only.

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Propagation and planting

Lophira alata is usually propagated by seed. The weight of 1000 seeds is about 1 kg. Under natural circumstances, fruits are disseminated by strong winds; with their wings they can cover distances of hundreds of meters before falling on the ground. The germination rate is high (80–95%), but the viability of seeds drops quickly because of the fairly rapid degradation of the seed oil. Four months after harvesting, the germination rate has fallen to about 25% and therefore seed has to be sown soon after collection. Soaking in water for several hours is recommended and the preferred sowing depth is 1–2 cm. Germination can be fairly rapid taking 9–16 days, but can also take longer, 18–30 days for normal, mono-embryonic seeds, and about 45 days for poly-embryonic seeds, which are fairly common in Lophira alata.
In the nursery plantlets are spaced at 10–15 cm, and have to be exposed to full sunlight soon. Small rodents are a threat because they dig up and eat the seeds. Seedlings form a taproot with many very fine lateral roots. Roots should be pruned frequently by shifting the containers to avoid that the taproot anchors itself in the soil and to stimulate the growth of lateral roots; this increases the transplantation success rate.
Transplanting into the field can be done when the plants are 18 months old, have a 40–100 cm long stem, a well-developed taproot and up to 15 leaves. However, transplanting younger plants is possible. Sometimes 15–20 cm tall wildlings are collected from natural forest in June and these are planted out after 12 months. Direct planting of 30–35 cm tall wildlings results in a lower success rate. Precautions should be taken to protect the young plants as they are sensitive to sun-scald and can dry out quickly. However, they are tolerant of poor soils. Young plants respond well to initial light shade, e.g. by interplanting between rows of existing trees producing light shade. The canopy has to be opened as soon as the young plants are well established. Ring weeding is required to avoid competition for water, nutrients and light, and when plants are well established, more complete weedings can be carried out. Young plants are more sensitive to attack by stem borers when they are exposed to full light; such attacks are more frequent in the nursery than in forest. Planting out in swampy localities has failed because the taproot died quickly.
Lophira alata coppices well, but does not form root suckers, unlike Lophira lanceolata. In Côte d’Ivoire, in-vitro culture of embryos of Lophira alata using various growth media (gel, silicagel, liquid) have shown growth rates of 2–4 cm/day, while the average daily growth rate was 1–2 cm.

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Management

In coastal forest on alluvial soils Lophira alata is often abundant or even dominant among canopy trees. The density of trees with a bole diameter of more than 50 cm can reach an average of 3 trees/ha in dense forest, with a local variation ranging from 1 to 7 trees/ha. At the beginning of the 20^th century, densities as high as 10–20 trees/ha with a bole diameter of more than 50 cm have been recorded. In inland areas in Gabon, where dense forest and more open Marantaceae forest meet, the density of large trees (bole diameter ≥ 70 cm) and of small trees (bole diameter ≥ 10 cm) is higher in open forest (reaching 0.9 trees/ha and 27.4 trees/ha, respectively) than in nearby mixed dense forest where the density is almost too low to measure.
Typical Lophira alata forest is dominated by large trees forming a continuous canopy, a dense undergrowth and the nearly absence of lianas. The distribution of bole diameters over size classes forms a bell-shaped curve, indicating that the recruitment of young trees is not assured under the cover of adult trees. Measurements of regeneration in dense forest have shown that seedlings could be abundant, but that the vast majority of them were less than 50 cm tall, while very few plants reached a bole diameter of 10–15 cm, except where they were favoured by natural or man-made forest openings.
Coppicing for the production of firewood has been studied in Côte d’Ivoire; close planting (2 m × 2 m) and coppicing at stump level, leaving 1–2, fairly straight and vigorous shoots per stump, are recommended. In plantations for timber, the recommended density is 1100 stems/ha with a first thinning after 6–8 years reducing the density to 500–600 stems/ha. When interplanting in food crops, both the use of small seedlings, which farmers tend to remove while weeding, and large seedlings (more than 1 m tall), which do not strike well and are sensitive to wind, should be avoided.
In Cameroon and Gabon Lophira alata is self-pruning and trees drop branches regularly and without leaving visible scars. In Côte d’Ivoire, however, branches are dropped later, leaving large protuberances as visible scars and leading to a more sinuous bole.
Management of plantations is generally limited to removal of small lianas and saplings of other trees. Some foresters recommend planting at high density or the establishment of mixed plantations in order to increase the establishment rate and delay branching.

Diseases and pests

Leaves of Lophira alata are often attacked by an insect, causing 4–5 cm large galls on both surfaces. In plantations, attacks have been reported by larvae of an unidentified insect destroying the apical bud, leading to premature branching of young stems.

Harvesting

Trees of Lophira alata are felled at any time of the year as long as forest tracks are passable. Harvesting methods with little destructive impact are sometimes used, but so far only in concessions engaged in certified production. The logs are too heavy to transport by water and have to be moved by truck.
Seeds are collected from under mother trees; bark for medicinal purposes is normally harvested with a machete.

Yield

Lophira alata is one of the species most widely exploited for its wood, in particular in Cameroon, Equatorial Guinea and Gabon. The minimum harvestable bole diameter is set at 60 cm in Cameroon and Côte d’Ivoire, 70 cm in Ghana, and 80 cm in Gabon and Liberia. An average harvestable tree yields 8–13.5 m³ of bole wood volume. A tradable volume of 24.2 m³ has been measured for a tree in Nigeria. Volume tables to estimate the productivity of populations are available for several countries.
Plantations in Abobo (Côte d’Ivoire), interplanted in food crops, have produced after 21 years a dominant canopy of Lophira alata at a height of 25–30 m with an average bole diameter of 20–25 cm. In Akila (Nigeria), a pure stand of Lophira alata had the following characteristics after 12 years: 1428 trees/ha, average bole diameter 10.6 cm, average height 16.3 m with the dominant canopy layer at 18.3 m, volume of the stand 67.5 m³ with an average annual growth rate of 5.6 m³/ha. Results obtained in a plantation in Sibang (Gabon) after 64 years indicate an estimated log volume of 252 m³/ha, or an annual growth rate of 3.9 m³/ha.

Handling after harvest

The removal of sapwood is recommended to reduce the risk of damage by rot-causing agents. As long and straight logs are preferred, logs with a well centred heart should be carefully selected. Boles of azobé with an excentric heart contain invisible tension wood and compression wood, which cause deformations as soon as pieces are lumbered. Although the blunting effect is small, sawing of azobé requires adapted and sufficiently powerful equipment.

Genetic resources and breeding

Because of the very high level of exploitation in its area of distribution, IUCN has classified Lophira alata as ‘vulnerable’. In almost the whole coastal region of Cameroon, populations have become seriously degraded or have disappeared completely as a result of human activities (urbanisation, road construction, food and industrial crop production, silvicultural exploitation). The species is now partially protected in a number of conservation areas, but this is insufficient to ascertain that regeneration in exploited areas will be adequate to meet future demands. In addition, the effects of the systematic exploitation of the best trees on the genetic diversity are not known. It may well have led to negative genetic selection, as the only seed trees remaining were the most poorly formed ones.

Breeding

No selection programmes for azobé currently exist.

Prospects

In West Africa and Cameroon, primary populations of Lophira alata have disappeared since the 1950s and 1960s. They are now almost everywhere degraded or have disappeared completely as a result of human activity in the coastal regions. It must be assumed that erosion of the genetic diversity has occurred. However, its characteristics as a pioneer species favour natural regeneration of azobé and offer possibilities for the establishment of plantations for timber production that can meet future demands. The sensitivity of azobé to competition from other pioneer species should be carefully kept in mind. Effective measures for its regeneration should be taken systematically by the logging companies that exploit azobé. Research on the genetics of the species and the selection of plants which are more tolerant to drought could favour its plantation on a wider scale.

Major references

• Aubréville, A., 1959. La flore forestière de la Côte d’Ivoire. Deuxième édition révisée. Tome deuxième. Publication No 15. Centre Technique Forestier Tropical, Nogent-sur-Marne, France. 341 pp.
• Bouquet, A., 1969. Féticheurs et médecines traditionnelles du Congo (Brazzaville). Mémoires ORSTOM No 36. Office de la Recherche Scientifique et Technique Outre-Mer. Paris, France. 282 pp.
• Burkill, H.M., 1997. The useful plants of West Tropical Africa. 2nd Edition. Volume 4, Families M–R. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 969 pp.
• CTFT (Centre Technique Forestier Tropical), 1976. Azobé. Bois et Forêts des Tropiques 170: 35–50.
• 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.
• Letouzey, R., 1957. La forêt à Lophira alata de la zone littorale camerounaise. Bois et Forêts des Tropiques 53: 9–20.
• Letouzey, R., 1985. Notice de la carte phytogéographique du Cameroun au 1 : 500 000. IRA, Yaoundé, Cameroun & Institut de la Carte Internationale de la Végétation, Toulouse, France. 5 volumes, 240 pp.
• Palla, F., Louppe, D. & Doumenge, C., 2002. Azobé. Fiche Technique, Projet Forafri, Libreville, Gabon. 4 pp.
• Takahashi, A., 1978. Compilation of data on the mechanical properties of foreign woods (part 3) Africa. Shimane University, Matsue, Japan. 248 pp.
• 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.

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Other references

• ATIBT (Association Technique Internationale des Bois Tropicaux), 1986. Tropical timber atlas: Part 1 – Africa. ATIBT, Paris, France. 208 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.
• CE-FAO, 1999. Données statistiques des produits forestiers non-ligneux du Cameroun. Rapport technique. Programme de partenariat CE-FAO (1998–2001), Rome, Italy. 36 pp.
• Cordiez, F., 2000. Etude des mécanismes de régénération naturelle de l’agba (Gossweilerodendron balsamiferum Harms), l’azobé (Lophira alata Banks ex Gaertn.f.), le movingui (Distemonanthus benthamianus Baill.) et l’ozigo (Dacryodes buettneri (Engl.) H.J. Lam) au Gabon. FUSAGx, Gembloux, Belgium. 88 pp.
• CTFT (Centre Technique Forestier Tropical), 1954. Monographie de Azobé, Lophira procera A. Chev. CTFT, Nogent-sur-Marne, France. 80 pp.
• de la Mensbruge, G., 1966. La germination et les plantules des essences arborées de la forêt dense humide de la Côte d’Ivoire. Centre Technique Forestier Tropical, Nogent-sur-Marne, France. 389 pp.
• Ewola Tih, A., Ghogomu Tih, R., Sondengam, B.L., Caux, C. & Bodo, B., 2006. Minor biflavonoids from Lophira alata leaves. Journal of Natural Products 69: 1206–1208.
• Eyog Matig, O., Ndoye, O., Kengue, J. & Awono, A. (Editors), 2006. Les fruitiers forestiers comestibles du Cameroun. IPGRI Regional Office for West and Central Africa, Cotonou, Benin. 204 pp.
• Hall, J.B. & Swaine, M.D., 1981. Distribution and ecology of vascular plants in a tropical rain forest: forest vegetation of Ghana. W. Junk Publishers, the Hague, Netherlands. 383 pp.
• Hecketsweiler, P., 1992. Phénologie et saisonnalité en forêt gabonaise. L’exemple de quelques espèces ligneuses. 2 volumes. Thèse dee doctorat, Université de Montpellier 2, France. 414 pp.
• Jiofack, T., Fokunang, C., Guedje, N., Kemeuze, V., Fongnzossie, E., Nkongmeneck, B.A., Mapongmetsem, P.M. & Tsabang, N., 2010. Ethnobotanical uses of medicinal plants of two ethnoecological regions of Cameroon. International Journal of Medicine & Medical Science 2(3): 60–79.
• Keay, R.W.J., 1954. Ochnaceae. In: Keay, R.W.J. (Editor). Flora of West Tropical Africa. Volume 1, part 1. 2nd Edition. Crown Agents for Oversea Governments and Administrations, London, United Kingdom. pp. 221–232.
• Martineau, M., 1933. Note sur les Lophira de forêt et de savane. Revue de Botanique Appliquée 13: 467–469.
• Méniaud, J., 1950. L’azobé et ses utilisations. Bois et Forêts des Tropiques 15: 261–266.
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Author(s)

•	C. Doumenge CIRAD, Campus International de Baillarguet TA-C-105/D, F-34398 Montpellier Cédex 5, France
•	V.O. Séné Herbier National du Cameroun (Institut de Recherche Agricole pour le Développement IRAD), B.P. 1601, 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 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. & Séné, V.O., 2012. Lophira alata Banks ex C.F.Gaertn. [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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