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Habitat Differentiation of Lauraceae K Blackwell Science, LtdOxford, UK ERE Ecological Research 0912-38142003 Ecological Society of Japan 181January 2003 539 Habitat differentiation of Lauraceae K. Sri-Ngernyuang et al. 10.1046/j.0912-3814.2002.00539.x Original Article114BEES SGML Ecological Research (2003) 18, 1–14 Habitat differentiation of Lauraceae species in a tropical lower montane forest in northern Thailand KRIANGSAK SRI-NGERNYUANG,1* MAMORU KANZAKI,2 TAKASHI MIZUNO,1 HIDEYUKI NOGUCHI,1 SAKHAN TEEJUNTUK,3 CHETTHA SUNGPALEE,3 MASATOSHI HARA,4 TAKUO YAMAKURA,1 PONGSAK SAHUNALU,3 PRICHA DHANMANONDA3 AND SARAYUDH BUNYAVEJCHEWIN5 1Graduate School of Science, Osaka City University, Sumiyoshi, Osaka 558-8585, Japan, 2Graduate School of Agriculture, Kyoto University, Kyoto 606-8502, Japan, 3Faculty of Forestry, Kasetsart University, Bangkok 10900, Thailand, 4Natural History Museum and Institute, Chiba 260-8682, Japan and 5Silvicultural Research Division, Royal Forest Department, Bangkok 10900, Thailand Dependency on topographical habitat was examined for Lauraceae tree species in a lower montane forest using a large-scale research plot established at Doi Inthanon National Park, northern Thailand. Twenty species of 10 genera of Lauraceae were recorded in a 7.5-ha part of the plot; Lauraceae accounted for 18% of the total basal area. Lauraceae was the most species-rich and most abundant family in the plot. In a cluster analysis based on the matrix of spatial associations between species, two clusters were recognized. Members of one cluster seemed to associate with lower-elevation habitats, and members of the other associated with habitats on ridges. By subdividing the study plot into 20 m ¥ 20 m squares, a discriminant analysis could be applied to the presence–absence data for the 17 species that had sufficient population density. The predictor variables used were the relative elevation, slope inclination, slope direction (transformed to deviation from SSW) and slope convexity for each of the squares. The discriminant models were tested statistically by applying the random shift technique. The models were significant for 11 of the species (65% of the species examined) and were associated with the topographical condition of the habitat. Stepwise selection of the predictor variables for these 11 species revealed that relative elevation and slope convexity were the most important factors for predicting the presence or absence of the Lauraceae species. Both these variables were considered to indicate the hydrological condition of the habitat. Key words: discriminant analysis; Doi Inthanon; large-scale plot; randomization test; topographical habitat. INTRODUCTION Oak-laurel forests are the dominant vegetation type in the mountains of tropical Asia, from the Fagaceae and Lauraceae are more abundant and Himalayas to New Guinea, and are closely related more likely to reach the top layer of forest canopies to the temperate evergreen oak forests of East Asia in the tropical montane zone of South-East Asia (Ohsawa 1991; Tagawa 1995). In South-East Asia, than in the lowland forests of the same region. The studies of montane forest zonation have been con- term ‘oak-laurel forest’ has been used for this veg- ducted on mountains in the Philippines (Aragones etation type (Kochummem 1989; Tagawa 1995). 1996; Buot & Okitsu 1998; Pipoly & Madulid 1998), Indonesia (Yamada 1977; Ohsawa et al. 1985; Abdulhadi et al. 1998), Malaysia (Proctor et al. 1988; Kitayama 1992; Nakashizuka et al. *Author to whom correspondence should be addressed. Present address: Faculty of Agricultural Pro- 1992), Brunei (Pendry & Proctor 1997) and duction, Maejo University, Chiang Mai 50290, Thai- Thailand (Ogawa et al. 1961; Robbins & land. Email: [email protected] Smitinand 1966; Santisuk 1988; Maxwell 1995; Received 22 August 2001. Maxwell et al. 1997). Geographic pattern of mon- Accepted 17 July 2002. tane forests were also conducted for this region 2 K. Sri-Ngernyuang et al. (Ohsawa 1991, 1995; Tagawa 1995). Studies using Recently, a critical problem associated with the large-scale research plots have been carried out in statistical test of habitat dependency has been lowland forests since the 1980s (Condit 1995; identified. Most statistical tests require that the Ashton 1998). By contrast, there have been few sampling units be independent. However, many similar intensive studies of the biodiversity, struc- studies have illustrated that most tree species dis- ture, and dynamics of montane forests. We estab- play a patchy distribution pattern (e.g. Hubbell lished a large-scale, 15-ha research plot in a 1979; Condit et al. 2000), and the abundances of well-developed montane forest in northern different tree species in continuously arranged Thailand, to fill this gap in montane forest studies. sampling units are spatially autocorrelated (Rossi Many factors play important roles in determin- et al. 1992; Roxburgh & Chesson 1998; Webb & ing the distribution of tree species in a plant com- Peart 2000). This means that the assumption of munity. Grubb (1977) categorized these factors randomness is violated, and thus traditional statis- into four component niches: habitat, life-form, tical tests cannot be applied (Roxburgh & Chesson phenological and regeneration niches. Large-scale 1998; Webb & Peart 2000). To avoid this prob- research plots are suitable for studying the associ- lem, we applied the ‘torus randomization’ or ‘ran- ation between habitat and plant distribution dom shift’ technique to test the significance of because such plots usually include various topo- habitat dependency in this study. graphical units and the trees can be enumerated throughout the plot, as shown by Hubbel and Foster (1986), Itoh (1995), Yamada et al. (1997) and Svenning (1999). In these studies, topograph- STUDY SITE ical preference was observed in at least some of the component species. Topographic segregation may A 15-ha research plot was established in a well- contribute to the maintenance of species richness developed, well-protected, lower montane rain for- in the communities. In this study, we focused on est, located at approximately 1700 m altitude in topographical habitat differentiation among Doi Inthanon National Park (18˚31¢13≤ N, Lauraceae, as one of a series of intensive studies on 98˚29¢44≤ E). Doi Inthanon is the highest moun- this family. Lauraceae is the most abundant and tain in Thailand (2565 m altitude), and lies in species-rich family and includes members ranging the southernmost extension of the Himalayan from upper-canopy species to under-canopy species foothills. in this plot (Hara et al. 2002). Therefore, a sub- Several authors have classified the vegetation of sample using Lauraceae trees may represent varia- Thailand (e.g. Ogawa et al. 1961; Smitinand & tion in the whole plant community along Nalamphun 1967). Santisuk (1988) summarized topographic gradients. those studies and divided the forest vegetation of Many studies on the habitat dependency of trees northern Thailand into two forest zones: a lowland use subjectively defined habitat types, such as zone (150–1000 m altitude) and a montane zone valley-bottom and hilltop habitat (e.g. Hubbell & (over 1000 m altitude). The montane-zone forest Foster 1986; Yamada et al. 1997; Svenning 1999). is subdivided into lower and upper montane for- The definition of habitat type is always somewhat ests. The transition between the lower and upper arbitrary, making comparisons between studies by montane forests lies at approximately 1800 m on different authors difficult. For this reason, we used Doi Inthanon, and coincides roughly with the the formula proposed by Yamakura et al. (1995) to lower limit of the cloud zone. numerically characterize topography. The use of Doi Inthanon is formed by a huge granite mass this method allows unequivocal between-site com- and three major rock types, gneiss, granite and parisons. Another advantage is that the method limestone, are found in the park (Pendleton can distinguish the relative importance of ele- 1962). Gneiss covers approximately half of the ments that characterize microhabitat topography, area of the park. The summit area of Doi Intha- such as slope inclination and direction. The non, including the 15-ha research plot, consists of method also allows a wider choice of statistical Pre-Cambrian gneissic rock (Faculty of Forestry methods for studies of habitat dependency. 1992; TDRI 1997). The substratum rocks on Doi Habitat differentiation of Lauraceae 3 Inthanon have produced a coarse, sandy loam soil tion of each square was obtained by averaging the (Pendleton 1962). The soil of the montane forest elevations of its four corners. The surface relief of has a thick, coarse-textured B horizon, exceeding the slope was expressed using the index of slope 60 cm in depth (Faculty of Forestry 1992). convexity proposed by Yamakura et al. (1995). The The climate of northern Thailand is strongly index was obtained by subtracting the mean ele- influenced by the monsoon, and three distinct sea- vation of 12 posts, set at 20-m intervals along the sons are recognizable. The rainy season is from sides of the 60 m ¥ 60 m square surrounding each June to October. A cool, dry period follows from focal 20 m ¥ 20 m square, from the elevation of the November to February, and a hot, dry season is focal square. Therefore, a positive value of the from March to May. Data from the Royal Thai Air index means that the slope relief is convex. Slope Force radar base, located at the summit (2565 m), convexity could not be calculated for the marginal are available from 1982 to 1999. The annual mean squares, because these squares lacked outer daily minimum and maximum temperatures are squares. 7.2°C and 18.6°C, respectively. The annual rain- This study was conducted using one half of the fall is 2279 mm per year. Data collected from 1993 15-ha plot. All woody plants (excluding climb- to 1999 at the Royal Project Doi Inthanon Station ers, bamboos and palms) of at least 1 cm d.b.h. (at 1300 m) show that the annual mean minimum (stem diameter 140 cm above the ground) in the and maximum temperatures are 16.1°C and 7.5-ha area (250 m ¥ 300 m) were counted and 25.7°C, respectively.
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