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Dipterocarp Forest in Lambir Hills National

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Dipterocarp Forest in Lambir Hills National Journal of Tropical Forest Science 14(3): 379-400 (2002) 379 FLORISTIC AND STRUCTURAL DIVERSITY OF MIXED DIPTEROCARP FORES LAMBIN TI R HILLS NATIONAL PARK, SARAWAK, MALAYSIA H. S. Lee, Forest Department, 93660 Kuching, Sarawak, Malaysia S. J. Davies*, Arnold Arboretum, Harvard University, Divinity22 Avenue, Cambridge MA02138, USA. E-mail: sdavies@oeb. harvard, edu J. V. LaFrankie, Center for Tropical Forest Science, Smithsonian Tropical Research Institute, Bukit Timah Road, Singapore S. Tan, Forest Department, 93660 Kuching, Sarawak, Malaysia . YamakuraT Itoh. ,A , Osaka City University, Sumiyoshi-ku, Osaka 558, Japan T. Ohkubo Utsunomiya National University, Utsunomiya 321-8505, Japan & Ashto. S . P n Arnold Arboretum, Harvard University, Divinity22 Avenue, Cambridge MA02138,USA Received March 2001_______________________________________________ LEE, H. S., DAVIES, S. J., LAFRANKIE, J. V., TAN, S., YAMAKURA, T., ITOH, A., OHKUBO, T. & ASHTON, P. S. 2002. Floristic and structural diversity of mixed dipterocarp fores Lambin i t r Hills National Park, Sarawak, Malaysia 52-hA . a permanent forest ploestablishes twa Lambin di r Hills National Park Sarawak, Malaysia to begin the long-term study of factors controlling the origin and maintenance of tree diversity. Stand structure and floristic composition of the plot are described. * Author for correspondence. 0 38 Journal of Tropical Forest Science 14(3): 379-400 (2002) Lambirwas found to have the highest diversity of trees anywhere on earth. In the 52-ha plot there were 356501 trees with a total basal area of 2252 m2, comprising 1173 tree species in 286 genera and 81 families. The Euphorbiaceae (125 species) and the Dipterocarpaceae (87 species) were the mostspecies-rich families. The Dipterocarpaceae dominate basaforese e dth th f l o t treesare e wit 16Burseraceaed e % th a%f han 42 Th .o , Anacardiacea Euphorbiacead ean e wer nexe eth t most important largee treeth n si plot. mose Shoreath s t importanwa t genus wit highesspeciee 3 h5 th d san t basal area and stem number. As with other Asian tropical forests there were many speciose gener plote th gener 1 ;n 2 a i >d a1ha 2 species addition I . Shorea,o nt Dipterocarpusd an Dryobalanops were important basal area contributor Dryobalanopsd san secone th s dwa most abundant genus. Dryobalanops aromatica and Dipterocarpus globosus were the most important canopy trees. Fordia splendidissima was the most abundant understorey tree importann A . ha 2 t5 findine ichange nth th s gi floristin ei c compositio stand nan d structure acros 52-he sth a plo relation i t soitopographid o nt an l c variation. Keywords: Basal are aBorne- odensit- yDipterocarpacea- eresourc- e heterogeneity - species richnes - stre e species diversit y- tropica l rain forest LEE, H. S., DAVIES, S. J., LAFRANKIE, J. V., TAN, S., YAMAKURA, T., ITOH, A., OHKUBO, T. & ASHTON, P. S. 2002. Kepelbagaiaii flora dan struktur hutan dipterokarpa campur di Tainan Negara Bukit Lambir, Sarawak, Malaysia. Petak hutan kekal seluas 52 ha ditubuhkan di Taman Negara Bukit Lambir, Sarawak, Malaysia bagi memulakan kajian jangka panjang tentang faktor yang mengawal asal dan penyenggaraan kepelbagaian pokok. Struktur diria kandungan nda n flora petak ini diterangkan. Lambir mempunyai kepelbagaian pokok yang tertinggi di dunia. Di dalam petak 52 ha terdapat 356 501 pokok dengan jumlah luas pangkal 2252 m2, terdirn da i daripada!173 spesies pokok dala gener6 m28 a da famili1 n8 . Euphorbiaceae (125 spesies Dipterocarpacean )da spesies7 e(8 ) merupakan famili yang mempunyai paling banyak spesies. Dipterocarpaceae menaungi hutan sebanyak 42% daripada luas pangkal dan 16% daripada pokok. Burseraceae, Anacardiaceae dan Euphorbiaceae merupakan jenis pokok besar yang kedua pentingny i dalaad m petak tersebut. Shorea ialah genus terpenting yang mempunyai 53 spesies dengan luas pangkal dan bilangan batang yang tertinggi. Seperti spesies hutan tropika Asia yang lain, terdapat banyak genera specios i dalaed m petak ini, iait gener1 u2 a mempunya ispesies 2 >1 . Di samping Shorea, Dipterocarpus Dryobalanopsn da merupaka a spesiendu s penting yang menyumbangkan luas pangkal. Dryobalanops ialah genus yang kedua banyaknya. Dryobalanops aromatica n Dipterocarpusda globosus merupakan pokok kanopi yang terpenting. Fordia splendidissima merupakan pokok tingkat bawah yang paling banyak di dalam petak 52 ha. Penemuan penting ialah terdapat perubahan dalam kandungan flora dan struktur dirian di sepanjang petak 52 ha apabila dikaitkan dengan variasi tanah dan topografi. Introduction Lowland tropical rain fores amons i t mose gth t diverse vegetatio worlde th n ni (Whitmore 1984, Richards 1996). The diversity of trees in these forests increases wit hdeclina climatin ei c seasonality, wit mose hth t diverse forests occurrinn gi areas without a significant annual dry period (Clinebell et al. 1995, Gaston 2000). Among lowland tropical forests of the aseasonal zone there is nevertheless a wide rang leveln ei tref so e diversity. Species richnes thesn si e forests maybe influenced by variatio soin ni l nutrient status (Ashto Haln& l 1992, Duivenvoorden 1996, Journal of Tropical Forest Science 14(3): 379-400 (2002) 381 Sollins 1998) probabilite th , short-terf yo m drought (Walsh 1996) abilite th , f yo certain species to become dominant and have a disproportionate access to limited resources (Connel Lowmal& n 1989, Har al.t e t 1989, Newber al.t ye 1997) wels a , l as historical and biogeographic factors (GunatillekeSc Ashton 1987). Resolving the relative influenc f theseo e different factor tropican so l forest diversit centraa s yi l concer tropicaf no l forest ecology (Givnish 1999). western I n Malesia, lowland forests wit highese hth t tree species diversite yar thought to occur on soils of intermediate nutrient status (Baillie et al. 1987, Ashton 1995, Davies & Becker 1996). Several studies have found lowest tree diversity in edaphically extreme sites r examplefo , , forest ultrabasin o s c soils (Procto. al t e r 1988), limestone-derived soils (Proctor et al. 1983), or white sand podzols (Brunig 1974, Procto l 1983a t re , Davie Beckes& r 1996) thesl al n eI . case soiw lslo nutrient availability appears to strongly influence the floristic composition of the forests, although other resources, particularly soil water availabilitye b co-limi y y ,ma ma r to more importan affectinn i t g forest composition (Walsh 1996). Some evidence suggests that forest soiln o f higso s h nutrient availabilit alsy oyma have lower species richness than forests on intermediate soils. Ashton and Hall (1992) found lower species richness on deep basalt-derived soils at Bukit Mersing in Sarawak, than on shallower, more nutrient-poor soils at Bukit Lambir. Furthermore, in Brunei, species richness was higher on sandy humult ultisols at Andulau than on more fertile clay-rich ultisol t Ladasa n (Davie Beckes& r 1996), althoug soio hn l nutrient analyses were conducte thesn di e sites. Stand structure also varies significantly among lowland tropical forests (Clark 1999a). eal t western I . n Malesia, mixed dipterocarp forest red-yellon so w podzolic soil generalle sar y talle staturn ri havd ean e much greater above-ground biomass than heath (kerangas) forests on white-sand podzols (Ashton & Hall 1992). In Gunung Mulu, Sarawak, mixed dipterocarp forest on sandstone-derived soil had 42% greater above-ground biomass than forest over limestongreate% 28 d r ean biomass than heath fores n acio t d podzols (Procto . 1983)al t e r . Heath forests typically also have a more even canopy with fewer emergents, greater densities smallese f treeo th n si t size classes predominanca d an , f specieeo s with small sclerophyllous leaves (Richards 1936, Browne 1952, Becke . 1999)al t re . Within these forest formations ther evidencs ei e that variatio stann ni d structure floristid an c compositio relates ni environmentao dt l gradients analysin A .8 3 f so small plot heatn si h fores Sarawan i t Bruned kan i revealed strong differencen i s floristic composition related to topography and soil properties (Brunig 1974, Newbery 1991). In mixed dipterocarp forests over the broad class of red-yellow podzolic soils, there is also substantial variation in structure and floristics related to variation in topography, drainage, soil clay content (and, by inference, soil nutrients) and geography (Austin et al 1972, Baillie et al. 1987, Ashton & Hall 1992). Ashto d Halnan l (1992) exampler ,fo , found that ste largese mth densitiet tbu l al n si size classes were lower on nutrient-rich soils at Bukit Mersing than on nutrient-poor soil t Bukisa t Lambir. Furthermore, comparison floristif so c similarity withie nth dominant family, Dipterocarpaceae, among mixed dipterocarp forests in Sarawak and Brunei found greatest similarity of dipterocarp composition for plots on 382 Journal of Tropical Forest Science 14(3): 379-400 (2002) similar soils and parent materials (Davies & Becker 1996). However, most studies of environmental correlates of forest composition in western Malesia have used small sample plots (usuall t accoun e effec no ) thath f locayo ha r o d t> 1 fo t l environmental variation on forest structure and composition. Also, in many cases, studies comparing forest differenn so t soil froe sar m geographically distant areas and, therefore, soil fertility differences may covary with other factors such as rainfall. In this paper, we describe floristic composition and stand structure of the mixed dipterocarp forest at Lambir Hills National Park, Sarawak, East Malaysia. Following similar projects initiated in Barro Colorado Island, Panama (Hubbell & Foster 1983) and Pasoh Forest Reserve in Peninsular Malaysia (Kochummen & LaFrankie 1990, Manokaran & LaFrankie 1990), this study encompasses a large fores facilitato t ) t ha are 2 e a(5 demographi c analyse significana f so t proportiof no tree th e species enablo t d e,an analyse importance th f so spatiaf eo l heterogeneity of resource availability for tree species and forest community distribution patterns (Condit 1995, Ashton et al.
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