We examined relationships between mortality rate, relative growth rate (RGR), and spatial patterns of three growth stages (small, medium, and large trees) for 11 dipterocarp species in the Pasoh 50-ha plot. Mortality rates for these species tended to be positively correlated with RGRs, although the correlation was significant only at the small-tree stage. Seven species with high growth and mortality rates exhibited peaks in spatial aggregation at small distances (<100 m) in small trees, but this aggregation disappeared in medium and large trees. In contrast, the other four species with low growth and mortality rates aggregated at large distances (>200 m) throughout the three growth stages in all but one species. Negative associations between different growth stages were observed only for the high-mortality species, suggesting density-dependent mortality. The high-mortality species showed habitat associations with topography, soil type, and the forest regeneration phase after gap formation, whereas the three low-mortality species only had associations with the forest regeneration phase. A randomization procedure revealed that these habitat associations explained little of their spatial aggregation. Our results suggest that the growth strategy has a large effect on the structuring of the spatial distribution of tree species through mortality processes.
Dispersal-assembly theories of species coexistence posit that environmental factors play no role in explaining community diversity and structure. Dispersal-assembly theories shed light on some aspects of community structure such as species-area and species-abundance relationships. However, species' environmental associations also affect these measures of community structure. Measurements of species' niche breadth and overlap address this influence. Using a new continuous measure of niche and a dispersal-assembly null model that maintains species' niche breadth and aggregation, we tested two hypotheses assessing the effects of habitat heterogeneity on the ability of dispersal-assembly theories to explain community niche structure. We found that in both homogenous and heterogeneous environments dispersal-assembly theories cannot fully explain observed niche structure. The performance of the dispersal-assembly null models was particularly poor in heterogeneous environments. These results indicate that non-dispersal based mechanisms are in part responsible for observed community structure and measures of community structure which include species' environmental associations should be used to test theories of species diversity.
The impacts of global change on tropical forests remain poorly understood. We examined changes in tree growth rates over the past two decades for all species occurring in large (50-ha) forest dynamics plots in Panama and Malaysia. Stem growth rates declined significantly at both forests regardless of initial size or organizational level (species, community or stand). Decreasing growth rates were widespread, occurring in 24-71% of species at Barro Colorado Island, Panama (BCI) and in 58-95% of species at Pasoh, Malaysia (depending on the sizes of stems included). Changes in growth were not consistently associated with initial growth rate, adult stature, or wood density. Changes in growth were significantly associated with regional climate changes: at both sites growth was negatively correlated with annual mean daily minimum temperatures, and at BCI growth was positively correlated with annual precipitation and number of rainfree days (a measure of relative insolation). While the underlying cause(s) of decelerating growth is still unresolved, these patterns strongly contradict the hypothesized pantropical increase in tree growth rates caused by carbon fertilization. Decelerating tree growth will have important economic and environmental implications.
The response of tropical forests to global climate variability and change remains poorly understood. Results from long-term studies of permanent forest plots have reported different, and in some cases opposing trends in tropical forest dynamics. In this study, we examined changes in tree growth rates at four long-term permanent tropical forest research plots in relation to variation in solar radiation, temperature and precipitation. Temporal variation in the stand-level growth rates measured at five-year intervals was found to be positively correlated with variation in incoming solar radiation and negatively related to temporal variation in night-time temperatures. Taken alone, neither solar radiation variability nor the effects of night-time temperatures can account for the observed temporal variation in tree growth rates across sites, but when considered together, these two climate variables account for most of the observed temporal variability in tree growth rates. Further analysis indicates that the stand-level response is primarily driven by the responses of smaller-sized trees (less than 20 cm in diameter). The combined temperature and radiation responses identified in this study provide a potential explanation for the conflicting patterns in tree growth rates found in previous studies.
The responses of tropical forests to global anthropogenic disturbances remain poorly understood. Above-ground woody biomass in some tropical forest plots has increased over the past several decades, potentially reflecting a widespread response to increased resource availability, for example, due to elevated atmospheric CO2 and/or nutrient deposition. However, previous studies of biomass dynamics have not accounted for natural patterns of disturbance and gap phase regeneration, making it difficult to quantify the importance of environmental changes. Using spatially explicit census data from large (50 ha) inventory plots, we investigated the influence of gap phase processes on the biomass dynamics of four 'old-growth' tropical forests (Barro Colorado Island (BCI), Panama; Pasoh and Lambir, Malaysia; and Huai Kha Khaeng (HKK), Thailand). We show that biomass increases were gradual and concentrated in earlier-phase forest patches, while biomass losses were generally of greater magnitude but concentrated in rarer later-phase patches. We then estimate the rate of biomass change at each site independent of gap phase dynamics using reduced major axis regressions and ANCOVA tests. Above-ground woody biomass increased significantly at Pasoh (+0.72% yr(-1)) and decreased at HKK (-0.56% yr(-1)) independent of changes in gap phase but remained stable at both BCI and Lambir. We conclude that gap phase processes play an important role in the biomass dynamics of tropical forests, and that quantifying the role of gap phase processes will help improve our understanding of the factors driving changes in forest biomass as well as their place in the global carbon budget.
While the importance of local-scale habitat niches in shaping tree species turnover along environmental gradients in tropical forests is well appreciated, relatively little is known about the influence of phylogenetic signal in species' habitat niches in shaping local community structure. We used detailed maps of the soil resource and topographic variation within eight 24-50 ha tropical forest plots combined with species phylogenies created from the APG III phylogeny to examine how phylogenetic beta diversity (indicating the degree of phylogenetic similarity of two communities) was related to environmental gradients within tropical tree communities. Using distance-based redundancy analysis we found that phylogenetic beta diversity, expressed as either nearest neighbor distance or mean pairwise distance, was significantly related to both soil and topographic variation in all study sites. In general, more phylogenetic beta diversity within a forest plot was explained by environmental variables this was expressed as nearest neighbor distance versus mean pairwise distance (3.0-10.3 % and 0.4-8.8 % of variation explained among plots, respectively), and more variation was explained by soil resource variables than topographic variables using either phylogenetic beta diversity metric. We also found that patterns of phylogenetic beta diversity expressed as nearest neighbor distance were consistent with previously observed patterns of niche similarity among congeneric species pairs in these plots. These results indicate the importance of phylogenetic signal in local habitat niches in shaping the phylogenetic structure of tropical tree communities, especially at the level of close phylogenetic neighbors, where similarity in habitat niches is most strongly preserved.