Comparative Biology of Tropical Trees: a Perspective from Pasoh

Published Date

pp 171-194

Title 

Comparative Biology of Tropical Trees: a Perspective from Pasoh

• Author 

• Sean C. Thomas

Abstract

From the late 1960s, tree species at Pasoh Forest Reserve (Pasoh FR) have been the focus of numerous comparative studies. Research to date has examined spatial distributions and edaphic preferences, reproductive phenology, vertical stratification, and interactions with pollinators and seed predators. Recent studies have also begun to address physiological and morphological bases for ecological variation. Pasoh FR is dominated by groups of trees that are phylogenetically closely related. As many as 47 species within a single genus (Eugenia) occur in sympatry at the site. Detailed studies of several of these speciose groups indicate that described morphospecies correspond to reproductivelyisolated taxa. Closely related species commonly differ in terms of reproductive timing, microhabitat preference, and vertical stratum at maturity. Comparative studies have also emphasized several bases for ecological “niche differentiation” at Pasoh, both within and among phylogenetic groups. Tree species at the site show pronounced differences in terms of vertical stratification, and relative to local patterns of water flow, soil parent material, and light requirements for growth. Recent theoretical studies suggest that niche differentiation may not be necessary to maintain high species richness in tropical forest communities. How, then, should one interpret the large ecological differences found among tree species at Pasoh FR? The existence of large interspecific differences in such characteristics as demographic rate parameters and size at reproduction indicates that one central assumption of “null” models-that species are equivalent-is simply incorrect. Models that fully incorporate interspecific ecological differences as well as dispersal limitation and other central elements of null models of tropical forest communities are needed to further our theoretical understanding of tropical tree diversity, and to better predict tropical forest responses to human and natural disturbance. Comparative studies are essential to parameterize and test such models, and also serve an important role in providing basic “silvics” information essential to forest management and conservation in the region.

References

1. Allbrook, R. F. (1973) The soils of Pasoh Forest Reserve, Negeri Sembilan. Malay. For. 36: 22–33.
2. Appanah, S. (1990) Plant-pollinator interactions in Malaysian rain forests. In Bawa, K. S. & Hadley, M. (eds). Reproductive Ecology of Tropical Forest Plants. UNESCO, Paris, pp.85–102.
3. Appanah, S. & Weinland, G (1993) Planting quality timber trees in Malaysia. Malaysian Forest Record No. 38. Forest Research Institute Malaysia, Kepong, Malaysia.
4. Appanah, S., Gentry, A. H. & LaFrankie, J. V. (1993) Liana diversity and species richness of Malaysian rain forests. J. Trop. For. Sci. 6: 116–123.
5. Appanah, S., Weinland, G, Bossel, H. & Krieger, H. (1990) Are tropical rain forests nonrenewable? An enquiry through modelling. J. Trop. For. Sci. 2: 331–348.
6. Ashton, P. S. (1969) Speciation among tropical forest trees: some deductions in the light of recent evidence. Biol. J. Linn. Soc. 1: 155–196.CrossRef
7. Ashton, P. S. (1976) Mixed dipterocarp forest and its variation with habitat in the Malayan lowlands: a re-evaluation at Pasoh. Malay. For. 39: 56–72.
8. Ashton, P. S. (1977) A contribution of rainforest research to evolutionary theory. Ann. Mo. Bot. Gard. 64: 694–705.CrossRef
9. Ashton, P. S. (1988) Dipterocarp biology as a window to the understanding of tropical forest structure. Annu. Rev. Ecol. Syst. 19: 347–370.CrossRef
10. Ashton, P. S., Givnish, T. J. & Appanah, S. (1988) Staggered flowering in the Dipterocarpaceae: New insights into floral induction and the evolution of mast fruiting in the aseasonal tropics. Am. Nat. 132: 44–66.CrossRef
11. Basset, Y., Aberlenc, H. & Delvare, G (1992) Abundance and stratification of foliage arthopods in a lowland rain-forest of Cameroon. Ecol. Entomol. 17: 310–318.CrossRef
12. Beccaloni, G (1997) Vertical stratification of ithomiine butterfly (Nymphalidae: Ithomiinae) mimicry complexes: the relationship between adult flight height and larval host-plant height. Biol. J. Linn. Soc. 62: 313–341.
13. Becker, P. & Wong, M. (1985) Seed dispersal, seed predation, and juvenile mortality ofAglaia sp. (Meliaceae) in lowland dipterocarp rainforest. Biotropica 17: 230–237.CrossRef
14. Bergh, J. P., van den & Braakhekke, W. G (1978) Coexistence of plant species by niche differentiation. In Freysen, A. H. J. & Woldendorp, J. W. (eds). Structure and functioning of plant populations. North-Holland, Amsterdam, pp.125–138.
15. Bossel, H. & Krieger, H. (1991) Simulation model of natural tropical forest dynamics. Ecol. Model. 59: 37–71.CrossRef
16. Bossel, H. & Krieger, H. (1994) Simulation of multi-species tropical forest dynamics using a vertically and horizontally structured model. For. Ecol. Manage. 69: 123–144.CrossRef
17. Brokaw, N. & Busing, R. T. (2000) Niche versus chance and tree diversity in forest gaps. Trend. Ecol. Evol. 15: 183–188.CrossRef
18. Burns, R. M. & Honkala, B. H. (1990) Silvics of North American Trees. vol. 2. Hardwoods. US Department of Agriculture, Washington, DC, USA.
19. Chan, H. T. & Appanah, S. (1980) Reproductive biology of some Malaysian dipterocarps. I. Flowering biology. Malay. For. 43: 132–143.
20. Chesson, P. L. & Warner, R. R. (1981) Environmental variability promotes coexistence in lottery competitive systems. Am. Nat. 117: 923–943.CrossRef
21. Clark, D. A. & Clark, D. B. (1992) Life history diversity of canopy and emergent trees in a neotropical rain forest. Ecol. Monogr. 62: 315–344.CrossRef
22. Clark, D. B., Clark, D.A. & Read, J. M. (1998) Edaphic variation and the mesoscale distribution of tree species in a neotropical rain forest. J. Ecol. 86: 101–112.CrossRef
23. Condit R, Ashton, P. S., Manokaran, N., LaFrankie, J. V., Hubbell, S. P.. & Foster, R. B. (1999) Dynamics of the forest communities at Pasoh and Barro Colorado: comparing two 50 ha plots. Philos. Trans. R. Soc. London Ser. B 354: 1739–1748.CrossRef
24. Condit, R., Ashton, P., Baker, P., Bunyavejchewin, S., Gunatilleke, S., Gunatilleke, N., Hubbell, S., Foster, R., Itoh, A., LaFrankie, J. V., Lee, H., Losos, E., Manokaran, N., Sukumar, R. & Yamakura T. (2000) Spatial patterns in the distribution of tropical tree species. Science 288: 1414–1418.PubMedCrossRef
25. Connell, J. H. (1971) On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees. In Den Boer, P. J. & Gradwell, G. (eds). Dynamics of Populations. PUDOC, Wageningen, the Netherlands, pp.298–312
26. Curran L. M., Caniago, I., Paoli, G D., Astianti, D., Kusneti, M., Leighton, M., Nirarita, C. E. & Haeruman, H. (1999) Impact of El Nino and logging on canopy tree recruitment in Borneo. Science 286: 2184–2188.PubMedCrossRef
27. Davies, S. J., Palmiotto, P. A., Ashton, P. S., Lee, H. S. & LaFrankie, J. V. (1999) Comparative ecology of 11 sympatric species of Macaranga in Borneo: tree distribution in relation to horizontal and vertical resource heterogeneity. J. Ecol. 86: 662–673.CrossRef
28. Denslow, J. S. (1980) Gap partitioning among tropical rainforest trees. Biotropica 12: 47–55.CrossRef
29. De Oliveira, A. & Mori, S. (1999) A central Amazonian terra firme forest. I. High tree species richness on poor soils. Biodiversity Conserv. 8: 1219–1244.CrossRef
30. Dobzhansky, T. (1950) Evolution in the tropics. Am. Sci. 38: 209–221.
31. Gentry, A. H. (1982) Neotropical floristic diversity: phytogeogrpahical connections between Central and South America, Pleistocene climatic fluctuations or an accident of Andean orogeny? Ann. Mo. Bot. Gard. 69: 557–593.CrossRef
32. Gentry, A. H. (1988a) Tree species richness of upper Amazonian forests. Proc. Natl. Acad. Sci. USA 85: 159–159.
33. Gentry, A. H. (1988b) Changes in plant community diversity and floristic composition on environmental and geographical gradients. Ann. Mo. Bot. Gard. 75: 1–34.CrossRef
34. Givnish, T. J. (1988) Adaptation to sun and shade: a whole plant perspective. Aust. J. Plant Physiol. 15: 63–92.CrossRef
35. Grubb, P. J. (1977) The maintenance of species richness in plant communities: the importance of the regeneration niche. Biol. Rev. 52: 107–145.CrossRef
36. Grubb, P. J. (1996) Rainforest dynamics: the need for new paradigms. In Edwards, D. S. (ed). Tropical rainforest research-current issues. Kluwer, Amsterdam, the Netherlands, pp.215–233.CrossRef
37. Ha, C. O., Sands, V. E., Soepadmo, E. & Jong, K. (1988) Reproductive patterns of selected understorey trees in the Malaysian rain forest: the apomictic species. Bot. J. Linn. Soc. 97: 317–331.CrossRef
38. He, F., Legendre, P., Bellehumeur, C. & LaFrankie, J. V. (1994) Diversity pattern and spatial scale: A study of a tropical rain forest of Malaysia. Environ. Ecol. Stat. 1: 265–286.CrossRef
39. He, F., Legendre, P. & LaFrankie, J. V. (1996) Spatial pattern of diversity in a tropical rain forest in Malaysia. J. Biogeogr. 23: 57–74.CrossRef
40. He, F., Legendre, P. & LaFrankie, J. V. (1997) Distribution patterns of tree species in a Malaysian tropical rain forest. J. Veg. Sci. 8: 105–114.CrossRef
41. Hubbell, S. P. (1995) Towards a theory of biodiversity and biogeography on continuous landscapes. In Carmichael, G R., Folk, G E. & Schnoor, J. L. (eds). Preparing for Global Change: A Midwestern Perspective. SPB Academic Publishing, Amsterdam, pp.171–199.
42. Hubbell, S. P. (1997) A unified theory of biogeography and relative species abundance and its application to tropical rain forests and coral reefs. Coral Reefs 16: S9–S21.CrossRef
43. Hubbell, S. P. (2001) The unified neutral theory of biodiversity and biogeography. Princeton University Press, Princeton, NJ, USA, 448pp.
44. Hubbell, S. P. & Foster, R. B. (1986) Biology, chance, and history and the structure of tropical rainforest tree communities. In Diamond, J. & Case, T. J. (eds). Community Ecology. Harper & Row, New York, pp.314–329.
45. Huth, A. & Ditzer, T. (2001) Long-term impacts of logging in a tropical rain forest- a simulation study. For. Ecol. Manage. 142: 33–51.CrossRef
46. Ickes, K. (2001) Density of native wild pigs (Sus scrofa) in a lowland Dipterocarp rain forest of Peninsular Malaysia. Biotropica 33: 682–690.
47. Ickes, K., De Walt, S. J. & Appanah, S. (2001) Effects of native pigs (Sus scrofa) on woody understorey vegetation in a Malaysian lowland rain forest. J. Trop. Ecol. 17: 191–206.CrossRef
48. Janzen, D. H. (1970) Herbivores and the number of tree species in tropical forests. Am. Nat. 104: 501–529.CrossRef
49. Janzen, D. H. (1974) Tropical black-water rivers, animals, and mast-fruiting in the Dipterocarpaceae. Biotropica 6: 69–103.CrossRef
50. Kato, M. (1996) Plant-pollinator interactions in the understory of a lowland mixed dipterocarp forest in Sarawak. Am. J. Bot. 83: 732–743.CrossRef
51. Kato, M., Inoue, T., Hamid, A., Nagamitsu, T., Merdek, M., Nona, A., Itino, T., Yamane, S. & Yumoto, T. (1995) Seasonality and vertical structure of light-attracted insect communities in a dipterocarp forest in Sarawak. Res. Popul. Ecol. 37: 59–79.CrossRef
52. Kaur, A., Ha, C. O., Hong, K., Sands, V. E., Chan, H. T., Soepadmo, E. & Ashton, P. S. (1978) Apomixis may be widespread among trees of the climax rain forest. Nature 271: 75–88.CrossRef
53. Kaur, A., Jong, K., Sands, V. E. & Soepadmo, E. (1986) Cytoembryology of some Malaysian dipterocarps, with some evidence of apomixis. Bot. J. Linn. Soc. 92: 75–88.CrossRef
54. Klopfer, P. & MacArthur, R. (1961) On the causes of tropical species diversity: niche overlap. Am. Nat. 95: 223–226.CrossRef
55. Kobe, R. K. (1999) Light gradient partitioning among tropical tree species through differential seedling mortality and growth. Ecology 80: 187–201.CrossRef
56. Kochummen, K. M. (1997) Tree Flora of Pasoh Forest. Kuala Lumpur, Malaysia: Forest Research Institute Malaysia, 462pp.
57. Kochummen, K. M., LaFrankie, J. V. & Manokaran, N. (1991) Floristic composition of Pasoh Forest Reserve, a lowland rain forest in Peninsular Malaysia. J. Trop. For. Sci. 3: 1–13.
58. Kohyama, T. (1993) Size-structured tree populations in gap-dynamic forest — the forest architecture hypothesis for the stable coexistence of species. J. Ecol. 81: 131–144.CrossRef
59. LaFrankie, J. V. & Chan, H. T. (1991) Confirmation of sequential flowering in Shorea (Dipterocarpaceae). Biotropica 23: 200–203.CrossRef
60. Lee, P. C. (1968) Studies on Dryobalanops aromatica. Ph.D. diss., Univ. Malaya.
61. Leigh, E. G, Wright, S. J., Herre, E. A. & Putz, F. E. (1993) The decline of tree diversity on newly isolated tropical islands — a test of a null hypothesis and some implications. Evol. Ecol. 7: 76–102.CrossRef
62. Liu, J. & Ashton, P. S. (1998) FORMOSAIC: an individual-based spatially explicit model for simulating forest dynamics in landscape mosaics. Ecol. Model. 106: 177–200.CrossRef
63. Liu, J. & Ashton, P. S. (1999) Simulating effects of landscape context and timber harvesting on tree species richness of a tropical forest. Ecol. Appl. 9: 186–201.CrossRef
64. Liu, J., Ickes, K., Ashton, P. S., LaFrankie, J. V. & Manokaran, N. (1999) Spatial and temporal impacts of adjacent areas on the dynamics of species diversity in a primary forest. In Mladenoff, D. J. & Baker, W. L. (eds). Spatial modeling of forest landscape change: approaches and applications. Cambridge University Press, Cambridge, UK, pp.42–69.
65. Manokaran, N., LaFrankie, J. V., Kochummen, K. M., Quah, E. S., Klahn, J. E., Ashton, P. S. & Hubbell, S. P. (1990) Methodology for the fifty-hectare research plot at Pasoh Forest Reserve. Research Pamphlet No. 104. Forest Research Institute Malaysia. Kepong, Malaysia, 69pp.
66. Manokaran, N., Kassim, A. R., Hassan, A., Quah, E. S. & Chong, P. F. (1992a) Short-term population dynamics of dipterocarp trees in a lowland rain forest in Peninsular Malaysia. J. Trop. For. Sci. 5: 97–112.
67. Manokaran, N., LaFrankie, J. V., Kotchummen, K. M., Quah, E. S., Klahn, J. E., Ashton, P. S. & Hubbell, S. P. (1992b) Stand tables and distribution of species in the fifty-hectare research plot at Pasoh Forest Reserve. FRIM Research Data Series 1. Forest Research Institute Malaysia, Kepong, Malaysia, 454pp.
68. Manokaran, N. & Swaine, M. D. (1994) Population dynamics of trees in dipterocarp forests of Peninsular Malaysia. Malayan Forest Record No. 40, Forest Research Intitute Malaysia, Kepong, Malaysia, 173pp.
69. Moad, A. S. (1993) Dipterocarp sapling growth and understory light availability in tropical lowland forest, Malaysia. Ph.D. diss., Harvard Univ., 340pp.
70. Momose, K., Yumoto, T., Nagamitsu, T., Kato, M., Nagamasu, H., Sakai, S., Harrison, R. D., Itioka, T., Hamid, A. A. & Inoue, T. (1998) Pollination biology in a lowland dipterocarp forest in Sarawak, Malaysia. I. Characteristics of the plant-pollinator community in a lowland dipterocarp forest. Am. J. Bot. 85: 1477–1501.PubMedCrossRef
71. Newman, E. I. (1982) Niche separation and species diversity in terrestrial vegetation. In Newman, E. I. (ed.) The plant community as a working mechanism. Special Publications No. 1 of the British Ecological Society. Blackwell Scientific Publications, Oxford, pp.61–78.
72. Oikawa, T. (1985) Simulation of forest carbon dynamics based on a dry-matter production model. I. Fundamental model structure of a tropical rainforest ecosystem. Bot. Mag. Tokyo 98: 225–238.CrossRef
73. Okuda, T., Kachi, N., Yap, S. K. & Manokaran, N. (1995) Spatial pattern of adult trees and seedling survivorship in Pentaspadon motleyi in a lowland rain forest in Peninsular Malaysia. J. Trop. For. Sci. 7: 475–489.
74. Okuda, T., Kachi, N., Yap, S. K. & Manokaran, N. (1997) Tree distribution pattern and fate of juveniles in a lowland tropical rain forest- implications for regeneration and maintenance of species diversity. Plant Ecol. 131: 155–171.CrossRef
75. Pitman, N. C. A., Terborgh, J., Silman, M. R. & Nuez, P. (1999) Tree species distributions in an upper Amazonian forest. Ecology 80: 2651–2661.CrossRef
76. Poore, M. E. D. (1968) Studies in Malaysian rain forest. I. The forest on triassic sediments in Jengka Forest Reserve. J. Ecol. 56: 143–196.CrossRef
77. Reich, P. B. (2000) Do tall trees scale physiological heights? Trends Ecol. Evol. 15: 41–42.PubMedCrossRef
78. Richards, A. J. (1990a) Studies in Garcinia, dieocious tropical forest trees: agamospermy. Bot. J. Linn. Soc. 103: 233–350.CrossRef
79. Richards, A. J. (1990b) Studies in Garcinia, dioecious tropical forest trees: the origin of the mangosteen (G. mangostana L.). Bot. J. Linn. Soc. 103: 301–308.CrossRef
80. Richards, P. W. (1969) Speciation in the tropical rain forest and the concept of the niche. Biol. J. Linn. Soc. 1: 149–153.CrossRef
81. Richards, P. W. (1978) Pasoh in perspective. Malay. Nat. J. 30: 145–148.
82. Rogstad, S. H. (1989) The biosystematics and evolution of the Polyalthia hypoleuca species complex (Annonaceae) of Malesia. I. Systematics. J. Arnold Arboretum 70: 153–246.
83. Rogstad, S. H. (1990) The biosystematics and evolution of the Polyalthia hypoleuca species complex (Annonaceae) of Malesia. II. Comparative distributional ecology. J. Trop. Ecol. 6: 387–408.CrossRef
84. Rogstad, S. H. (1994) Biosystematics and evolution of the Polyalthia hypoleuca species complex (Annonaceae) of Malesia. III. Floral ontogeny and breeding systems. Am. J. Bot. 81: 145–154.CrossRef
85. Sakai, S., Momose, K., Yumoto, T., Kato, M. & Inoue, T. (1999) Beetle pollination of Shorea parvifolia (section Mutica, Dipterocarpaceae) in a general flowering period in Sarawak, Malaysia. Am. J. Bot. 86: 62–69.PubMedCrossRef
86. Sato, T., Saw, L. G. & Furukawa, A. (2000) A quantitative comparison of pteridophytes diversity in small scales among different climatic regions in eastern Asia. Tropics 9: 83–90.CrossRef
87. Schot, A. M. (1995) A synopsis of taxonomic changes in Aporosa Blume (Euphorbiaceae). Blumea 40: 449–460.
88. Stevens, P. F. (1990) Nomenclatural stability, taxonomic instinct, and flora writing- a recipe for disaster? In Baas, P., Kalkman, K. & Geesink, R. (eds). The plant diversity of Malesia. Kluwer Academic Publisher, the Netherlands, pp.387–410.CrossRef
89. Svenning, J. C. (2000) Small canopy gaps influence plant distributions in the rain forest understory. Biotropica 32: 252–261.
90. Tang, Y. H., Kachi, N., Fukuyama, A. & Awang, M. (1996) Light reduction by regional haze and its effect on simulated leaf photosynthesis in a tropical forest of Malaysia. For. Ecol. Manage. 89: 205–211.CrossRef
91. Thomas, S. C. (1993) Interspecific allometry in Malaysian rainforest trees. Ph.D. diss., Harvard Univ., 291pp.
92. Thomas, S. C. (1996a) Asymptotic height as a predictor of growth and allometric characteristics in Malaysian rain forest trees. Am. J. Bot. 83: 556–566.CrossRef
93. Thomas, S. C. (1996b) Relative size at reproductive onset in rain forest trees: a comparative analysis of 37 Malaysian species. Oikos 76: 145–154.CrossRef
94. Thomas, S. C. (1996c) Reproductive allometry in Malaysian rain forest trees: biomechanics vs. optimal allocation. Evol. Ecol. 10: 517–530.CrossRef
95. Thomas, S. C. (1997) Geographic parthenogenesis in a tropical rain forest tree. Am. J. Bot. 84: 1012–1015.PubMedCrossRef
96. Thomas, S. C. (2003) Ecological correlates of tree species persistence in tropical forest fragments. In Losos, E. C. & Leigh, E. G. Jr. (eds). Forest diversity and dynamism: Findings from a network of large-scale tropical forest plots. University of Chicago Press, Chicago (in press).
97. Thomas, S. C. & LaFrankie, J. V. (1993) Sex, size, and interyear variation in flowering among dioecious trees of the Malayan rain forest. Ecology 74: 1529–1537.CrossRef
98. Thomas, S. C. & Bazaz, F. A. (1999) Asymptotic height as a predictor of photosynthetic characteristics in Malaysian rain forest trees. Ecology 80: 1607–1622.CrossRef
99. Thomas, S. C. & Ickes, K. (1995) Ontogenetic changes in leaf size in Malaysian rain forest trees. Biotropica 27: 427–434.CrossRef
100. Tilman, D. & Pacala, S. W. (1993) The maintenance of species richness in plant communities. In Ricklefs, R. E. & Schluter, D. (eds). Species diversity in ecological communities: historical and geographical perspectives. University of Chicago Press, Chicago, USA, pp.13–25.
101. Toy, R. J. (1991) Interspecific flowering patterns in the Dipterocarpaceae in West Malaysia: implications for predator satiation. J. Trop. Ecol. 7: 49–57.CrossRef
102. Toy, R., Marshall, A. & Pong, T. (1991) Fruiting phenology and the survival of insect fruit predators-a case-study from the south-east Asian Dipterocarpaceae. Philos. Trans. R. Soc. London Ser. B 335: 417–423.CrossRef
103. Valencia, R. (1998) Preliminary comparisons between Yasuni and BCI plots. Inside CTFS Summer 1998: 3.
104. Valencia, R., Balslev, H. & Pazy Miñio, C. G (1994) High tree alpha-diversity in Amazonian Ecuador. Biodiversity Conserv. 3: 21–28.CrossRef
105. Vandermeer, J. (1996) Disturbance and neutral competition theory in rain forest dynamics. Ecol. Model. 85: 99–111.CrossRef
106. Van Steenis, C. G G J. (1969) Plant speciation in Malesia, with special reference to the theory of non-adaptive saltatory evolution. Biol. J. Linn. Soc. 1: 97–133.CrossRef
107. Wallace, A. R. (1878) Tropical nature and other essays. MacMillan, London, UK.CrossRef
108. Warner, R. R. & Chesson, P. L. (1985) Coexistence mediated by recruitment fluctuations: a field guide to the storage effect. Am. Nat. 125: 769–787.CrossRef
109. Webb, C. O. & Peart, D. R. (2000) Habitat associations of trees and seedlings in a Bornean rain forest. J. Ecol. 88: 464–478.CrossRef
110. Whitmore, T. C. (1984) Tropical rain forests of the far east (2nd ed.). Clarendon Press, Oxford.
111. Wills, C. & Condit, R. (1999) Similar non-random processes maintain diversity in two tropical rainforests. Proc. R. Soc. London Ser. B 266: 1445–1452.CrossRef
112. Wills, C., Condit, R., Foster, R. B. & Hubbell, S. P. (1997) Strong density- and diversity-related effects help to maintain tree species diversity in a neotropical forest. Proc. Natl. Acad. Sci. USA 94: 1252–1257.PubMedCrossRef
113. Wolseley, P., Ellis, L., Harrington, A. & Moncrieff, C. (1998) Epiphytic cryptogams at Pasoh Forest Reserve, Negri Sembilan, Malaysia- Quantitative and qualitative sampling in logged and unlogged plots. In Lee, S. S., Dan, Y. M., Gauld, I. D. & Bishop, J. (eds). Conservation, management and development of forest resources: proceedings of the Malaysia-United Kingdom Programme Workshop, 21–24 October 1996, Kuala Lumpur, Malaysia. Forest Research Institute Malaysia, Kepong, Malaysia, pp.61–83.
114. Wong, M. (1981) Impact of dipterocarp seedlings on the vegetative and reproductive characteristics of Labisia pumila (Myrsiniaceae) in the understory. Malay. For. 44: 370–376.
115. Wong, M. (1983) Understory phenology of the virgin and regenerating habitats in Pasoh forest reserve, Negeri Sembilan, West Malaysia. Malay. For. 46: 197–223.
116. Wong, Y. K. & Whitmore, T. C. (1970) On the influence of soil properties on species distribution in a Malayan lowland dipterocarp forest. Malay. For. 33: 42–54.
117. Yamada, T., Itoh, A., Kanzaki, M., Yamakura, T., Suzuki, E. & Ashton, P. S. (2000) Local and geographical distributions for a tropical tree genus, Scaphium (Sterculiaceae) in the Far East. Plant Ecol. 148: 23–30.CrossRef
118. Yap, S. K. (1976) The reproductive biology of some understorey fruit tree species in the lowland dipterocarp forest of West Malaysia. Ph.D. diss., Univ. Malaya, 146pp.
119. Yap, S. K. (1982) The phenology of some fruit tree species in a lowland dipterocarp forest. Malay. For. 45: 21–35.
120. Yap, S. K. & Chan, H. T. (1990) Phenological behavior of some Shorea species in Peninsular Malaysia. In Bawa, K. S. & Hadley, M. (eds). Reproductive Ecology of Tropical Forest Plants. UNESCO, Paris, pp.21–35.

For further details log on website :
http://link.springer.com/chapter/10.1007/978-4-431-67008-7_13