Akira Itoh

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.

Consequences of defaunation for a tropical tree community.

Hunting affects a considerably greater area of the tropical forest biome than deforestation and logging combined. Often even large remote protected areas are depleted of a substantial proportion of their vertebrate fauna. However, understanding of the long-term ecological consequences of defaunation in tropical forests remains poor. Using tree census data from a large-scale plot monitored over a 15-year period since the approximate onset of intense hunting, we provide a comprehensive assessment of the immediate consequences of defaunation for a tropical tree community. Our data strongly suggest that over-hunting has engendered pervasive changes in tree population spatial structure and dynamics, leading to a consistent decline in local tree diversity over time. However, we do not find any support for suggestions that over-hunting reduces above-ground biomass or biomass accumulation rate in this forest. To maintain critical ecosystem processes in tropical forests increased efforts are required to protect and restore wildlife populations.

Scale‐dependent relationships between tree species richness and ecosystem function in forests

1. The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity.

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.

Habitat associations of Sterculiaceae trees in a Bornean rain forest plot

Abstract Questions: 1. Are trees in a Bornean tropical rain forest associated with a particular habitat? 2. Does the strength of habitat association with the species-specific optimal habitat increase with tree size? Location: A 52-ha plot in a mixed dipterocarp forest in a heterogeneous landscape at the Lambir Hills National Park, Sarawak, East Malaysia. Methods: Ten species from the Sterculiaceae were chosen as representative of all species in the plot, on the assumption that competition among closely related species is more stringent than that among more distantly related taxa. Their habitat associations were tested using data from a 52-ha plot by a torus-translation test. Results: The torus-translation test showed that eight out of the ten species examined had significant association with at least one habitat. We could not find negative species-habitat associations for rare species, probably due to their small sample sizes. Among four species small trees were less strongly associated with habitat than large trees, implying competitive exclusion of trees in suboptimal habitats. The other four species showed the opposite pattern, possibly owing to the smaller sample size of large trees. A habitat had a maximum of three species with which it was significantly positively associated. Conclusions: For a species to survive in population equilibrium in a landscape, habitats in which ‘source’ subpopulations can be sustained without subsidy from adjacent habitats are essential. Competition is most severe among related species whose source subpopulations share the same habitat. On the evidence of source subpopulations identified by positive species-habitat association, species-habitat association reduces the number of confamilial competitors. Our results therefore indicate that edaphic niche specialization contributes to coexistence of species of Sterculiaceae in the plot, consistent with the expectations of equilibrium hypotheses. Nomenclature: Ashton (1980).

Rooting ability of cuttings relates to phylogeny, habitat preference and growth characteristics of tropical rainforest trees

Abstract The rooting ability of branch cuttings was evaluated for 100 tree species (including 41 families and 78 genera) collected in a tropical rainforest in Sarawak, Malaysia. Leafy cuttings of natural forest saplings were planted in a non-mist propagation system with IBA treatment. During the 6-month experiment, 66 species were rooted with an overall mean rooting percentage of 37.7% (range 0–100%). Species in the families Dipterocarpaceae and Lauraceae had a low rooting ability, whereas those in Euphorbiaceae, Rubiaceae, and Annonaceae had a high rooting ability. Differences in rooting ability were related to species-specific mature sizes, diameter growth rates and habitat preferences. Species of smaller mature sizes and faster diameter growth rates showed better rooting ability. Species whose forest saplings sprouted more vigorously after experimental felling rooted better than those that showed less vigorous sprouting. Species whose habitats were on lower elevations, concave slopes, and/or clay-rich soils rooted significantly better than those that preferred opposite habitats or habitat generalists that showed no significant habitat preference. The implications of these relations are discussed from the viewpoint of saplings’ adaptation to physical damage in their natural habitats.

Habitat divergence in sympatric Fagaceae tree species of a tropical montane forest in northern Thailand

Spatial distributions of many tropical trees are skewed to specific habitats, i.e. habitat specialization. However, habitats of specialist species must be divergent, i.e. habitat divergence, to coexist in a local community. When a pair of species specialize in the same habitat, i.e. habitat convergence, they could not coexist by way of habitat specialization. Thus, analyses of habitat divergence, in addition to habitat specialization, are necessary to discuss coexistence mechanisms ofsympatric species. In this study, the habitat specialization and habitat divergence along topographic gradients of eight sympatric tree species of the Fagaceae were studied in a 15-ha study plot in a tropical lower montane forest in northern Thailand. A statistical test with torus shift randomizations for 9 6 73 trees of Fagaceae revealed significantly biased distributions for all of the species, for at least one of the four topographic variables used: elevation, slope inclination, aspect and convexity. Slope convexity was the most critical topographic variable, along which all but one species had significantly skewed distributions. Out of 112 possible combinations of species pairs and topographic variables, 18 (16%) and two pairs (1.8%) showed significant habitat divergence and habitat convergence, respectively. The observed habitat divergence alone could not completely explain the coexistence of the eight species. There was a gradation in the habitat position of each species, with relatively large overlaps between species distributed in similar habitats, and small overlaps between species associated with contrasting habitats, respectively. The gradual changes in the habitats of the species suggested that dividing the species into a small number of distinct habitat groups, such as ridge and valley specialists, would not be straightforward.

Intra- and interspecific variation in wood density and fine-scale spatial distribution of stand-level wood density in a northern Thai tropical montane forest.

Tropicaltreewooddensityisoftenrelatedtootherspecies-specificfunctionaltraits,e.g.size,growthrateand mortality. We would therefore expect significant associations within tropical forests between the spatial distributions of stand-level wood density and micro-environments when interspecific variation in wood density is larger than intraspecific variation and when habitat-based species assembly is important in the forest. In this study, we used wood cores collected from 515 trees of 72 species in a 15-ha plot in northern Thailand to analyse intra- and interspecific variationinwooddensityandthespatialassociationofstand-levelwooddensity.Intraspecificvariationwaslowerthan interspecific variation (20% vs. 80% of the total variation), indicating that species-specific differences in wood density, rather than phenotypic plasticity, are the major source of variation in wood density at the study site. Wood density of individual species was significantly negatively related to maximum diameter, growth rate of sapling diameter and mortalityofsaplings.Stand-levelmeanwooddensitywassignificantlynegativelyrelatedtoelevation,slopeconvexity, saplinggrowthrateandsaplingmortality,andpositivelyrelatedtoslopeinclination.East-facingslopeshadsignificantly lower stand-level mean wood densities than west-facing slopes. We hypothesized that ridges and east-facing slopes in the study forest experience strong and frequent wind disturbance, and that this severe impact may lead to faster stand turnover, creating conditions that favour fast-growing species with low wood density.

Spatially aggregated fruiting in an emergent Bornean tree

We investigated the spatial distribution of fruiting in Dryobalanops aromatica (Dipterocarpaceae), a mast-seeding emergent tree that is found in the tropical rain forests of Borneo. Of 393 adults (diameter at breast height (dbh) > 30 cm) in a 52-ha area, 143 (36.4%) individuals fruited. A second-order spatial analysis, based on Ripleys K-function, revealed that fruiting trees were significantly (P < 0.05) aggregated with respect to the total population over distances of 15-115 m. Thus, the spatial distribution of fruiting trees was more aggregated than expected from the adult tree distribution, which itself was aggregated within the study plot. Logistic regression analysis showed that fruiting trees had a significantly greater dbh and were surrounded by more conspecific adults per ha than non-fruiting individuals. Moreover, fruiting trees were found at higher elevations, on steeper slopes, and on sandy rather than clayey soils. In contrast, annual diameter growth, adult density over areas < 1 ha, and slope convexity did not significantly affect fruiting probability. The ratio of seedling (20 cm ≤ h < 60 cm) and sapling (60 cm ≤ h < 300 cm) density to adult density was higher in sites at higher elevations, on steeper slopes, and on sandy soils than that for lower elevations, flat slopes and clayey soils. This suggests that the aggregation of fruiting trees occurred not only at the time of the study, but that it had occurred repeatedly in the past, at the high-elevation, steep-sloped, sandy sites. Thus, site conditions probably affect the dynamics and spatial structure of local populations through differences in fruiting frequency among trees experiencing different site conditions.