Hostname: page-component-78c5997874-xbtfd Total loading time: 0 Render date: 2024-11-19T10:45:43.052Z Has data issue: false hasContentIssue false

Holes and the sums of parts in Ghanaian forest: regeneration, scale and sustainable use

Published online by Cambridge University Press:  05 December 2011

W. D. Hawthorne
Affiliation:
7 Poplar Rd, Botley, OX29LAOxford forestry Institute, South Parks RD, Oxford OX1 3RB
Get access

Synopsis

The current state of Ghana's forest is summarised. Considerable changes have occurred in the last decade, since Hall & Swaine's account and classification, due mainly to fire and logging. The requirements and potential for sustainable forest use are explored through a summary of patterns of regeneration, and of local and national distribution of individual species.

Incisive indices of forest quality and condition are vital to good forest management. Various forest quality indices, summarising different properties of the plant community, are examined. These indices gloss over the statistically noisy behaviour of single species in small forest areas. The indices are: Forest Type – Hall & Swaine's forest ordination and classification; a Pioneer Index (PI) revealing the balance of ‘regeneration guilds’; a Genetic Heat Index (GHI), based mainly on the rarity value (Star rating) of all forest species, highlighting ‘hotspots’; and an Economic Index (EI) based on the concentration of common species (‘reddish Stars’) threatened by exploitation. Guild and Star are defined for all species and encapsulate trends of local and of global distribution and ecology. The national and local patterns and response to disturbance of the indices derived from the representation of these various guilds and stars are discussed.

Scale is crucial to all discussions. A strictly hierarchical model of forest ecology/biogeography is less suitable than a continuum-of-significant-scale, and non-hierarchical model. For instance, refugia are usually perceived as discrete biogeographical units. However, major biological ‘hotspots’, which are often described as refugia and attributed to Pleistocene climatic variation, differ only in position along a continuum of scale from mini-refugia as small as individual plants. The biogeographic Dahomey gap has much in common with a canopy gap, with scale as the main distinction.

There are conspicuous trends across Ghana's forests in the abundance of pioneer, rare or economic species. These differ in detail, but ‘hysteresis’ – the forest memory – and other factors related to the concept of refugia apply to all these aspects of forest quality. Major hotspot refugia are crucial to the national framework of biodiversity, but local refugia, between the size of individual plants and single forest blocks, are crucial to local regeneration and sustainable use, as they shape the probability cloud which defines the anatomy of and processes within each species' range. Short-term sustainable use depends on local refugia; longer-term sustainability requires maintenance of refugia on a wider range of scale.

The implications of these phenomena to forest management are discussed in conclusion. Forest health is a multi-scale, but particularly a broad-scale, phenomenon. Local processes like the regeneration of forest under canopy gaps, are subordinate to larger-scale patterns and not determined simply by a match between species physiology and gap dynamics or patterns in the physical environment. Success of a species in a certain landscape does not automatically imply the species can be successful in similar conditions in a different landscape elsewhere: the context of the landscape in terms of the broader mosaic is also important. Managers, whether of plantations or natural forest, need to monitor, plan, and protect indigenous species on all scales. Forest managers need also to be aware of and work with the ‘forest memory’ factor. Protective measures for rare or economically threatened species should be based on current refugia and, like them, be arranged on all scales from single trees to large forest blocks.

Researchers need to pay more attention to processes between the ecological and biogeographical, if they are to provide information for managers which has a useful synergy with existing types of data. Exploration is needed of the anatomy of the ‘probability clouds’ defining the statistics of dispersal and regeneration of rare or threatened species with respect to parent populations. What are the chances of a mahogany establishing at a point 500 metres from a mother tree? How is this statistic influenced by soil type? There is much to be learnt on scales between the canopy and the Dahomey Gap.

Type
Research Article
Copyright
Copyright © Royal Society of Edinburgh 1996

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Abercrombie, M., Hickman, C. J. & Johnson, M. L. 1966. ‘A dictionary of biology’. (5th edn). London: Penguin.Google Scholar
Adams, D. E. & Anderson, R. C. 1982. An inverse relationship between dominance and habitat breadth of trees in Illinois forest. American Midland Naturalist 107, 192–5.Google Scholar
Alder, D. 1992. Simple methods for calculating minimum diameter and sustainable yield in mixed tropical forest. In Miller, F.R. & Adam, K.L. (Eds.), Wise management of Tropical Forests. Proceedings of the Oxford Conference on tropical forests Oxford: Oxford Forestry Institute pp. 189200.Google Scholar
Alexandre, D. Y. 1977. Régéneration naturelle d'un arbre caracteristique de la forêt équatorielle de Côte d'lvoire: Turraeanthus africana Pellegr. Oeceologia Plantarum 12(3), 241–66.Google Scholar
Allen, J. F. H. & Hoekstrom, T. W. 1992. Towards a unified ecology. New York: Columbia University Press.Google Scholar
Aubréville, A. 1938. La forêt coloniale. Annales, Academie des Sciences Coloniales. 9, 1245.Google Scholar
Blackett, H. L. 1989. Inventory techniques. In Wong, J. L. (Ed) Seminar Proceedings, Forests Inventory Project 1989. Overseas Development Administration (UK) and Forestry Department, Ghana.Google Scholar
Borman, F. H. & Likens, G. E. 1979. Pattern and process in a forested ecosystem. New York: Springer.Google Scholar
Borota, J. 1991. Tropical forests. Some African and Asian case studies of composition and structure. Developments in Agricultural and Managed–Forest Ecology 22 Oxford: Elsevier.Google Scholar
Brown, J. H., 1984. On the relationship between abundance and distribution of species. American Naturalist. 124, 255–79.Google Scholar
Brown, N. 1993. The implications of climate and gap microclimate for seedling growth conditions in a Bornean lowland rain forest. Journal Tropical Ecology 9, 153–68.Google Scholar
Brown, N. & Whitmore, T. C. 1992. Do dipterocarp seedlings really partition tropical rain forest gaps? Philosophical Transactions of Royal Society of London B. 335, 369–78.Google Scholar
Budowski, G. 1965. Distribution of tropical American rain forest species in the light of successional processes. Turrialba 15, 40–2.Google Scholar
Canham, C. D. 1989. Different responses to gaps among shade-tolerant tree species. Ecology 70, 548–50.Google Scholar
Cantor, L. F. & Whitham, T. G. 1989. Importance of belowground herbivory: pocket gophers may limit aspen to rock outcrop refugia. Ecology 70, 962–70.Google Scholar
Carter, R. N. & Prince, S. D. 1981. Epidemic models used to explain biogeographical distributional limits. Nature 293, 644–5.Google Scholar
Clark, D. A. & Clark, D. B. 1992. Life history diversity of canopy and emergent trees in a neotropical forest. Ecological Monographs 62, 315–44.CrossRefGoogle Scholar
Connor, E. F. 1986. The role of Pleistocene forest refugia in the evolution and biogeography of tropical biotas. Tree 1, 165–8.Google Scholar
Diamond, A. W. & Hamilton, A. C. 1980. The distribution of forest passerine birds and Quaternary climatic change in tropical Africa. Journal of Zoology London 191, 379402.CrossRefGoogle Scholar
Doran, J. C., Turnbull, J. W., Boland, D. J., Gunn, B. V. 1983. Handbook on seeds of dry-zone acacias. Rome: FAO.Google Scholar
Endler, J. A. 1982. Pleistocene forest refuges: fact or fancy. In Prance, G. T. (Ed.) Biological diversification in the tropics, pp. 641–57 New York: Columbia Univ. Press.Google Scholar
Epp, G. A. 1987. The seed bank of Eupatorium odoratum along a successional gradient in a tropical rain forest in Ghana. Journal of Tropical Ecology 3, pp. 18. Accra, Ghana.Google Scholar
François, J. H. 1989. Overview of Ghana's forest policy, Forestry Commission Symposia Series 3.Google Scholar
Gaston, K. J. 1988. Patterns in the local and regional dynamics of moth populations. Oikos 53, 4957.Google Scholar
Gaston, K. J. 1991. How large is a species geographic range? Oikos 61, 434–8Google Scholar
Gaston, K. J. & Lawton, J. W. 1990. Effects of scale and habitat on the relationship between regional distribution and local abundance. Oikos 58, 329–55.CrossRefGoogle Scholar
Gentry, A. H. 1989. Speciation in tropical forests. In Holm-Nielsen, L. B., Nielson, I. C. & Basler, H. (Eds) Tropical forests: botanical dynamics, speciation and diversity, pp. 113–34. London: Academic Press.Google Scholar
Gentry, A. H. 1992. Tropical forest biodiversity: distributional patterns and their conservational significance. Oikos 63, 1928.Google Scholar
Gomez-Pompa, A., Whitmore, T. C. & Hadley, M. (Eds). 1991. Rainforest regeneration and management. Paris: UNESCO.Google Scholar
Grubb, P. J. 1977. The maintenance of species richness in plant communities: the importance of the regeneration niche. Biological reviews 52, 107–45.Google Scholar
Guillanmet, J. L. 1967. Recherches sur la végétation et la flora de la région du Bas-Carally (Côte d'lvoire). Paris: Off. Rech. Scient. Tech. Outre-mer.Google Scholar
Hall, J. B. & Swaine, M. D. 1976. Classification and ecology of closed-canopy forest in Ghana. Journal of Ecology 64, 913–51.Google Scholar
Hall, J. B. & Swaine, M. D. 1981. Distribution and ecology of vascular plants in a tropical rain forest. The Hague: W. Junk.Google Scholar
Hallé, F., Oldeman, R. A. A., Tomlinson, P. B., 1978. Tropical trees and forests. Berlin: Springer-Verlag.Google Scholar
Hamilton, A. C. 1974. ‘Distribution patterns of forest refugia in Uganda and their historical importance’. Vegetatio 29, 3135.Google Scholar
Hamilton, A. C. 1982. The Quaternary history of East African forests: its relevance to conservation. African Journal of Ecology 19, 16.Google Scholar
Hamilton, A. C., & Taylor, D. 1991. History of climate and forests in tropical Africa during the last 8 million years. Climatic Change 19, 6578.Google Scholar
Hamilton, A. C., & Taylor, D. 1992. Are hotspots in African forests quaternary refugia. Paper presented at Wildlife Conservation International conference on conservation of African forests. June 1992.Google Scholar
Hanski, I. 1982. Dynamics of regional distribution, the core and satellite species hypothesis. Oikos 38, 210–21.Google Scholar
Hawkins, C. P. & MacMahon, J. A. 1989. Guilds: the multiple meanings of a concept. Annual Review of Entomology 34, 423–51.Google Scholar
Hawthorne, W. D. 1984. Ecological and biogeographical patterns in the coastal forests of East Africa. D. Phil. Thesis. University of Oxford.Google Scholar
Hawthorne, W. D. 1988. Regeneration of Ghana's forests. Unpublished report to Ghana Forestry dept. & Overseas Development Admin. (Superseded by Hawthorne, 1995.)Google Scholar
Hawthorne, W. D. 1989 & 1990 – see Hawthorne 1993c and 1994.Google Scholar
Hawthorne, W. D. 1990. Field Guide to the Forest Trees of Ghana. Chatham, UK: Natural Resources Institute.Google Scholar
Hawthorne, W. D. 1992. Forestry, Dragons and Genetic Heat. Paper presented at seminar on conservation in Africa, Wildlife Conservation Internation. June, 1992.Google Scholar
Hawthorne, W. D. 1993a. East African coastal forest botany. In Lovett, J. C. & Wasser, S. K. (Eds) Biogeography and ecology of the rain forests of eastern Africa. Cambridge University Press, pp. 5799.Google Scholar
Hawthorne, W. D. 1993b. ‘Froggie manual’ (parts 1 and 2) and software. To be distributed with Hawthorne & Abu Juam 1995 (see below).Google Scholar
Hawthorne, W. D. 1993c. Regeneration after logging in Bia South GPR, Ghana. Natural Resouces Institute, Chatham. U. K. (First circulated as report to ODA/ Ghana Forestry Department in 1989).Google Scholar
Hawthorne, W. D. 1994. Fire damage and forest regeneration in Ghana. Natural Resouces Institute, Chatham. U. K. (Circulated as report to ODA/ Ghana Forestry Department in 1990).Google Scholar
Hawthorne, W. D. & Abu Juam, M. 1995 Forest protection in Ghana. Gland, Switzerland and Cambridge, UK: IUCN. (Previously (1993) circulated as unpublished, ODA reports.)Google Scholar
Hawthorne, W. D., 1995. Ecological profiles of Ghanaian forest trees. Tropical Forestry Papers 29. Oxford Forestry Institute, UK.Google Scholar
Hengeveld, R. 1990. Dynamic biogeography. Cambridge Studies in Ecology. Cambridge University Press.Google Scholar
Hengeveld, R. & Haeck, J. 1982. The distribution of abundance. 1. Measurements. Journal of Biogeography 9, 303–6.Google Scholar
Hepper, F. N. (Ed.) 19541972. Flora of West Tropical Africa (2nd Edn). London: HMSO.Google Scholar
Holling, C. S. 1992. Cross scale morphology, geometry and dynamics of ecosystems. Ecological Monographs 62, 447502.Google Scholar
Holm-Nielsen, L. B. Nielsen, I. C. & Baslev, H. (Eds). 1989. Tropical forests: botanical dynamics, speciation and diversity. London: Academic Press.Google Scholar
Howe, H. F. & Smallwood, J. 1982. Ecology of seed dispersal. Annual Review of Ecology and Systematics 13, 201–28.Google Scholar
Hubbell, S. P. 1980. Seed predation and the coexistence of tree species in tropical forests. Oikos 35, 214–29.Google Scholar
Janzen, D. H. 1974. Tropical Blackwater rivers, animals and mast fruiting by the Dipterocarpaceae. Biotropica 6, 69103.Google Scholar
Janzen, D. H. 1978. Seedling patterns in tropical trees. In Tomlinson, P. B. & Zimmermann, M. H., (Eds) Tropical trees as living systems. Cambridge: Cambridge University Press.Google Scholar
Johnson, R. A. 1981. Application of the guild concept to environmental impact analysis of terrestrial vegetation. Journal of Environmental Management 13, 205–22.Google Scholar
Jones, E. W. 1957. Report of Chlorophora. London: HMSO.Google Scholar
Kennedy, D. N. & Swaine, M. D., 1992. Germination and growth of colonising species in artificial gaps of different sizes in dipterocarp rain forest. Philosophical Transactions of the Royal Society of London B 335, 357–66.Google Scholar
Kerr, J. R. 1987. Biological monitoring and environmental impact assessment: a conceptual framework. Environmental Management 11, 249–56.Google Scholar
Kingdon, J. S. 1980. The role of visual signals and face patterns in African forest monkeys (gueonons) of the genus Cercopithecus. Transactions of the Zoological Society of London 35, 425–74.Google Scholar
Lavorel, S., Godner, R. H. & O'Neill, R. V. 1993. Analysis of patterns in hierarchically structured landscapes. Oikos 67, 521–8.Google Scholar
Lawson, G. W., Armstrong-Mensah, K. O. & Hall, J. B. 1970. A catena in tropical moist semi-deciduous forest near Kade, Ghana. Journal of Ecology 58, 371–98.Google Scholar
Lawton, J. H. 1992. There are not 10 million kinds of population dynamics. Oikos 63, 337–8.Google Scholar
Leonard, J. 1989. Revision du genre Martretia Beille (Euphorbiaceae) et la nouvelle tribu des Martretieae. Bulletin du Jardin Botanique National de Belgique 59, 333–49.Google Scholar
Lieberman, M. & Lieberman, D. 1989. Forests are not just swiss cheese: canopy stereogeometry of non-gaps in tropical forests. Ecology 70, 550–52.Google Scholar
Livingstone, D. A. 1975. Late Quaternary climatic change in Africa. Annual Review of Ecology and Systematics 6, 249–80.Google Scholar
Lovett, J. C. & Wasser, S. K. (Eds) 1993. Biogeography and ecology of the rainforests of eastern Africa, Cambridge University Press.CrossRefGoogle Scholar
Macarthur, R. H. & Wilson, E. O. 1967. The theory of island biogeography. Princeton: Princeton University Press.Google Scholar
Maginnis, S. 1994. Understorey condition scoring of Ghanian lowland tropical moist forest during stock survey: a technique for regulating the allowable cut in ecologically and structurally degraded production forest. Forest Ecology & Management 70, 8997.Google Scholar
Maley, J. 1991. The African rain forest vegetation and palaeoenvironments during late Quaternary. Climatic Change 19, 7998.Google Scholar
Maley, J., 1995. The African rain forest: principal patterns of vegetation from Upper Cretaceous to Quaternary. Proceedings of the Royal Society of Edinburgh 104B, 00–00.Google Scholar
Martin, C. 1991. The rainforests of West Africa. (Translated from German by Tsardaka, L.). Birkhauser Verlag Ag.Google Scholar
Meave, J., Kellman, M., Macdougall, A., & Rosales, J. 1991. Riparian habitats as tropical forest refugia. Global Ecology and Biogeography Letters 1, 6976.CrossRefGoogle Scholar
Novotny, V. 1991. The effect of habitat persistence on the relationship between geographic distribution and local abundance. Oikos 61, 431–3.Google Scholar
Oldeman, R. A. A. 1989. Dynamics in tropical rain forests. In Holm-Nielsen, L. B., Nielson, L. C. & Baslev, H. (Eds) Tropical forests: botanical dynamics, speciation and diversity, pp. 325 London: Acadmeic Press.Google Scholar
Oldeman, R. A. A. 1990. Forests: elements of silvology. New York: Springer-Verlag.Google Scholar
Oldeman, R. A. A. & Van Dijk, J. 1991. Diagnosis of the temperament of tropical rain forest trees. In Gomez-Pompa, A., Whitmore, T. C. & Hadley, M. (Eds) Rain forest regeneration and management, pp. 2162. Paris: UNESCO.Google Scholar
Palmer, M. W. 1989. Fractal geometry: a tool for describing spatial patterns of plant communities. Vegetatio 75, 91102.Google Scholar
Parren, M. P. E. 1991. Forest elephant (Loxodonta africana cyclotis Matschie) messanger-boy or bulldozer? The possible impact as the vegetation, with special reference to 41 tree species of Ghana. AV 90/51. Wageningen Agricultural University, The Netherlands.Google Scholar
Pielou, E. C. 1981. The usefulness of ecological models: a stock-taking. Quarterly Review of Biology 56, 1723.Google Scholar
Pigott, C. D. 1981. Nature of seed sterility and the natural regeneration of Tilia cordata near its northern limit in Finland. Annales Botanici Fennici 18, 255–63.Google Scholar
Popma, J., Bongers, F., Martinez-Ramos, M., & Veneklaas, E. 1988. Pioneer species distribution in treefall gaps in neotropical rainforest: a gap definition and its consequences. Journal of Tropical Ecology IV, 7788.Google Scholar
Raich, J. W., & Gong, W. K. 1990 Effects of canopy opening on tree seed germination in a Malaysian dipterocarp forest. Journal of Tropical Ecology 6, 203–17.Google Scholar
Reader, R. 1988. Using the guild concept in the assessment of tree harvesting effects on understorey herbs: a cautionary note. Environmental Management 12, 803–8.Google Scholar
Rodgers, W. A., Owen, C. F. & Homewood, K. M. 1982. Biogeography of East African forest mammals. Journal of Biogeography 9, 4154.Google Scholar
Ross, R. 1954. Ecological studies on the rain forest of southern Nigeria. III. The secondary succession in the Shasha forest reserve. Journal of Ecology 42, 259–82.CrossRefGoogle Scholar
Severinghaus, W. D. 1981. Guild theory development as a mechanism for assessing environmental impact. Environmental Management 5, 187–90.Google Scholar
Shugart, H. H. 1984. A theory of rain forest dynamics. New York: Springer-Verlag.Google Scholar
Shugart, H. H. & Urban, D. L. 1989. Factors affecting the relative abundances of tree species. In Grubb, P. J. & Whittaker, J. B. (Eds). Towards a more exact ecology, pp. 249–73. 30th Symposium of British Ecological Society. Oxford: Blackwells Scientific.Google Scholar
Simberloff, D. & Dayan, T. 1991. The guild concept and the structure of ecological communities. Annual Review of Ecology and Systematics 22, 115–43.Google Scholar
Smith, T. M. & Urban, D. C. 1988. Scale and resolution of forest structural pattern. Vegetatio 74, 143–50.Google Scholar
Sosef, M. S. M. 1994. Studies in Begoniaceae V: Refuge Begonias. Taxonomy, phylogeny & historical biogeography of Begonia sect. Loasibegonia and sect. Scutobegonia in relation to glacial forest refuges in Africa. Wageningen Agricultural University Papers; 94.1.Google Scholar
Swaine, M. D. 1983. Stilt roots and ephemeral germination sites. Biotropica 15, 240.Google Scholar
Swaine, M. D. 1989. Population Dynamics of tree species in tropical forests. In Holm-Nielsen, L. B., Nielsen, L. C. & Baslev, H. (Eds) Tropical forests: botanical dynamics, speciation and diversity, pp. 101–13. London: Academic Press.Google Scholar
Swaine, M. D., & Hall, J. B. 1981. The monospecific tropical forest of the Ghanaian endemic tree Talbotiella gentii. In Synge, H. (Ed.) The biological aspects of rare plant conservation, proceedings of an international conference, p. 35 London: Wiley.Google Scholar
Swaine, M. D. & Hall, J. B. 1983. Early succession on cleared forest land in Ghana. Journal of Ecology 71, 601–28.Google Scholar
Swaine, M. D. & Hall, J. B. 1988. The mosaic theory of forest regeneration and the determination of forest composition in Ghana. Journal of Tropical Ecology 4, 253–69.Google Scholar
Swaine, M. D. & Whitmore, T. C. 1988. On the definition of ecological species groups in tropical rain forests. Vegetatio 75, 81–6.Google Scholar
Swaine, M. D., Hall, J. B. & Alexander, I. J. 1987. Tree population dynamics at Kade, Ghana (1968–1982). Journal of Ecology 3, 331–46.Google Scholar
Swaine, M. D., Lieberman, D. & Hall, J. B. 1990. Structure and dynamics of a tropical dry forest in Ghana. Vegetatio 88, 3151.Google Scholar
Swaine, M. D., Hawthorne, W. D. & Orgle, T. C. 1992. The effects of fire exclusion on savanna vegetation at Kpong, Ghana. Biotropica 24, 166–72.Google Scholar
Thornton, I. W. B. 1983. Vicariance and dispersal: confrontation or compatibility?. Geojournal, 7, 557–64.Google Scholar
Turner, M. G. 1989. Landscape ecology: The effect of pattern on process. Annual Review Ecology and Systematics 20, 171–98.Google Scholar
Van Rompaey, R. S. A. R., 1993. Forest gradients in West Africa: a spatial gradient analysis. Doctoral thesis, Wageningen Agricultural University, The Netherlands, xxii + 142 pp. ISBN 90–5485–120–1.Google Scholar
Veblen, T. T. 1989. Tree regeneration responses to gaps along a transandean gradient. Ecology 70, 541–3.Google Scholar
Voorhoeve, A. G. 1965. Liberian high forest trees. Wageningen: Centre for Agricultural Publication & Documentation.Google Scholar
Watt, A. S., 1947. Pattern and process in the plant community. Journal of Ecology 35, 122.Google Scholar
White, F. 1971. The taxonomic and ecological basis of chorology. Mitteilungen der Botanischein Staatssamlung Munchen 10, 91112.Google Scholar
White, F. 1978. The taxonomy, ecology and chorology of African Ebenaceae. 1. The Guineo-Congolian species. Bulletin du Jardin Botanique National de Belgique 48, 245358.Google Scholar
White, F. 1979. The Guineo-Congolian region and its relationship to other phytochoria. Bulletin du Jardin Botanique National de Belgique 49, 1155.Google Scholar
White, F. 1983a. The vegetation of Africa. A descriptive memoir to accompany the UNESCO/AETFAT/UNSO vegetation map of Africa. Paris: UNESCO.Google Scholar
White, F. 1983b. Long distance dispersal and the origins of the Afromontane flora. Sonderbande zur Naturwiss enschaftlicher Verlag Hamburg 7, 87116.Google Scholar
Whitmore, T. C. 1978. Gaps in the forest canopy. In Tomlinson, P. B. & Zimmermann, M. H. (Eds) Tropical trees as living systems, pp. 639–55 Cambridge: Cambridge University Press.Google Scholar
Whitmore, T. C., 1989. Canopy gaps and the two major groups of forest trees. Ecology 70, 536–8.Google Scholar
Whitmore, T. C. 1991. Tropical rain forest dynamics and its implications for management. In Gomez-Pompa, A., Whitmore, T. C. & Hadley, M. Rain forest regeneration and managment, pp. 6776. Paris: UNESCO.Google Scholar
Woods, P. 1989. Effects of logging, drought, and fire on structure and composition of tropical forests in Sabah, Malaysia. Biotropica 21, 290–98.Google Scholar
Wyatt-Smith, J. (1950). Virgin Jungle Reserves. Malayan Forester 13, 4045.Google Scholar