James V. LaFrankie

Distribution patterns of tree species in a Malaysian tropical rain forest

Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regu- lar distribution. The spatial patterns of rare vs. common spe- cies, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.

The Importance of Demographic Niches to Tree Diversity

Most ecological hypotheses about species coexistence hinge on species differences, but quantifying trait differences across species in diverse communities is often unfeasible. We examined the variation of demographic traits using a global tropical forest data set covering 4500 species in 10 large-scale tree inventories. With a hierarchical Bayesian approach, we quantified the distribution of mortality and growth rates of all tree species at each site. This allowed us to test the prediction that demographic differences facilitate species richness, as suggested by the theory that a tradeoff between high growth and high survival allows species to coexist. Contrary to the prediction, the most diverse forests had the least demographic variation. Although demographic differences may foster coexistence, they do not explain any of the 16-fold variation in tree species richness observed across the tropics.

Assessing Evidence for a Pervasive Alteration in Tropical Tree Communities

In Amazonian tropical forests, recent studies have reported increases in aboveground biomass and in primary productivity, as well as shifts in plant species composition favouring fast-growing species over slow-growing ones. This pervasive alteration of mature tropical forests was attributed to global environmental change, such as an increase in atmospheric CO2 concentration, nutrient deposition, temperature, drought frequency, and/or irradiance. We used standardized, repeated measurements of over 2 million trees in ten large (16–52 ha each) forest plots on three continents to evaluate the generality of these findings across tropical forests. Aboveground biomass increased at seven of our ten plots, significantly so at four plots, and showed a large decrease at a single plot. Carbon accumulation pooled across sites was significant (+0.24 MgC ha−1 y−1, 95% confidence intervals [0.07, 0.39] MgC ha−1 y−1), but lower than reported previously for Amazonia. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. Over all ten plots, the fastest-growing quartile of species gained biomass (+0.33 [0.09, 0.55] % y−1) compared with the tree community as a whole (+0.15 % y−1); however, this significant trend was due to a single plot. Biomass of slow-growing species increased significantly when calculated over all plots (+0.21 [0.02, 0.37] % y−1), and in half of our plots when calculated individually. Our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Instead, they suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability. More long-term studies are necessary to clarify the contribution of global change to the functioning of tropical forests.

Importance of topography and soil texture in the spatial distribution of two sympatric dipterocarp trees in a Bornean rainforest

Relationships between spatial distributions and site conditions, namely topography and soil texture, were analyzed for two congeneric emergent trees, Dryobalanops aromatica and Dryobalanops lanceolata (Dipterocarpaceae), in a tropical rainforest in Sarawak, East Malaysia. A 52-ha permanent plot was divided into 1300 quadrats measuring 20 m × 20 m; for each Dryobalanops species, the number and total basal area of trees ≥1 cm in d.b.h. were compared among groups of quadrats with different site conditions. Because spatial distributions of both Dryobalanops and site-condition variables were aggregated, Monte-Carlo permutation tests were applied to analyze the relationships. Both single and multifactor statistical tests showed that the density and basal area distributions of the two species were significantly non-random in relation to soil texture and topographic variables. D. aromatica was significantly more abundant at higher elevations, in sandy soils, and on convex and steep slopes. In contrast, D. lanceolata preferred lower elevations and less sandy soils. In the study plot, there were very few sites (3 of 1150 quadrats tested) where the models of Hayashi’s method predicted the co-occurrence of the two species. These results suggest that between-species differences in habitat preferences are so large that they alone explain the spatially segregated distributions of these two species within the 52-ha study plot.

Diversity pattern and spatial scale: a study of a tropical rain forest of Malaysia

Scale is emerging as one of the critical problems in ecology because our perception of most ecological variables and processes depends upon the scale at which the variables are measured. A conclusion obtained at one scale may not be valid at another scale without sufficient knowledge of the scaling effect, which is also a source of misinterpretation for many ecological problems, such as the design of reserves in conservation biology.This paper attempts to study empirically how scaling may affect the spatial patterns of diversity (tree density, richness and Shannon diversity) that we may perceive in tropical forests, using as a test-case a 50 ha forest plot in Malaysia. The effect of scale on measurements of diversity patterns, the occurrence of rare species, the fractal dimension of diversity patterns, the spatial structure and the nearest-neighbour autocorrelation of diversity are addressed. The response of a variable to scale depends on the way it is measured and the way it is distributed in space.We conclude that, in general, the effect of scaling on measures of biological diversity is non-linear; heterogeneity increases with the size of the sampling units, and fine-scale information is lost at a broad scale. Our results should lead to a better understanding of how ecological variables and processes change over scale.

HABITAT PREFERENCES OF APOROSA IN TWO MALAYSIAN FORESTS: IMPLICATIONS FOR ABUNDANCE AND COEXISTENCE

Theories accounting for the maintenance of high tree diversity in tropical rain forests range from those proposing that tropical trees are highly co-evolved niche specialists, to those proposing that they are mostly generalist, undergoing random drift. We test these hypotheses at a meaningful, community-wide scale using data on the spatial patterns and habitat preferences of all species of Aporosa (Euphorbiaceae) growing on two large rain forest plots in Malaysia. Second-order spatial pattern analyses using a method based on Ripleys K function showed that Aporosa species formed spatially distinct assemblages, and a randomization procedure suggested that these assemblages were explained by biases in their distributions in relation to habitat types. Soil type, as determined by parent material, was an important determinant of habitat preferences, although topography and forest structure also accounted for some variation. We conclude that niche differentiation is an important mechanism contributing to the coexistence of Aporosa species at the community scale. However, spatial separation due to these differential habitat biases accounted for only a portion of the high species richness observed in this genus, so other mechanisms must also be sought to account fully for the maintenance of tropical tree species richness.

Soil-Related Floristic Variation in a Hyperdiverse Dipterocarp Forest

A 52 hectare permanent research plot was established in Lambir Hills National Park to enable long-term study of factors controlling the origin and maintenance of tree diversity. In this chapter we summarize some of our recent work on the relationships between floristic variation and edaphic heterogeneity in the Lambir forest. First, we provide a general description of the floristic composition of this hyperdiverse forest. Second, we use a detailed survey of soil chemistry to test whether floristic composition changes in relation to edaphic characteristics. We also assess the extent to which individual species have non-random distributions in the forest with respect to edaphic heterogeneity. Finally, to investigate the influence of habitat variation on floristic diversity we compare our results from the heterogeneous forest at Lambir Hills with a more homogeneous forest in Peninsular Malaysia.

Scale dependence of tree abundance and richness in a tropical rain forest, Malaysia

Abundance and richness are the two fundamental components of speciesdiversity. They represent two distinct types of variables of which the formerisadditive when aggregated across scales while the latter is nonadditive. Thisstudy investigated the changes in the spatial patterns of abundance andrichnessof tree species across multiple scales in a tropical rain forest of Malaysiaandtheir variations in different regions of the study area. The results showedthatfrom fine to coarse scales abundance had a gradual and systematic change inpattern, whereas the change in richness was much less predictable and a‘hotspot’ in richness at one scale may become a‘coldspot’ at another. The study also demonstrated that differentmeasures of diversity variation (e.g., variance and coefficient of variation)can result in different or even contradictory results which further complicatedthe interpretation of diversity patterns. Because of scale effect the commonlyused measure of species diversity in terms of unit area (e.g.,species/m2) is misleading and of little use in comparing speciesdiversitybetween different ecosystems. Extra care must be taken if management andconservation of species diversity have to be based on information gathered at asingle scale.

population dynamics of some tropical trees that yield non-timber forest products

The population biology ofAquilaria malaccensis, one source of gharu, and Cinnamomum mollissimum, one source of wild cinnamon, was studied in a 50 ha permanent plot of primary rain forest in Malaysia. Median diameter growth rates of 0.22 cm yr-1 and 0.1 cm yr-1 should not be prohibitive of economic exploitation, and suggest that the trees could be grown commercially. However, the natural densities were between 2 and 3 trees over 1 cm d.b.h. per ha, which is roughly the median for all trees in the plot, would preclude economic exploitation of these natural populations. The economics of harvesting natural populations is considered in a preliminary fashion by allowing favorable assumptions of quantity and quality of production. The price likely to be fetched from either a first time extraction (on the order of US

CONTRASTING STRUCTURE AND COMPOSITION OF THE UNDERSTORY IN SPECIES‐RICH TROPICAL RAIN FORESTS

10.00 per ha) or from sustained production (on the order of US