1. One of the primary goals in community ecology is to determine the relative importance of processes and mechanisms that control biodiversity. Here, we examined habitat-driven species assemblages ...and species distribution patterns as well as their temporal variations for three life stages of two censuses of a 25-ha mixed dipterocarp forest at Sinharaja (Sri Lanka). 2. Our general objective was to find out whether the species assemblages and associated habitat types changed with life stage, spatial scale and species attributes. We also analyse whether the habitat types were related to certain indicator species. Habitat types were determined with multivariate regression tree analyses driven by topographic variables. 3. We found species assemblages associated with five distinct habitat types that appeared consistently for all life stages of the two censuses. These habitats were related to ridge-valley gradients and a pronounced contrast in south-west versus north-east aspect. Habitat-driven structuring was weak at the recruit stage but strong in the juvenile and adult stages. The species assemblage variance explained by topographic variables for different life stages ranged between 10% for recruits and 23% for juveniles. 4. The species assemblages determined for different spatial scales (10, 20, 50 m) showed similar habitat partitioning, but the variance explained by the topographic variables increased in all life stages with spatial scale. This could be due to the homogenizing effect of topographic variables at the larger scales and unaccounted environmental variation at the smaller scales. The number of indicator species identified in the two censuses was higher in the juvenile stage than in the adult stage, and nearly all indicator species in the adult stage were also indicator species in the juvenile stage. 5. Synthesis. Our study showed that approximately 75% of the variance in local species composition is unexplained. This may be due to spatially structured processes such as dispersal limitation, unaccounted biotic and abiotic environmental variables, and stochastic effects, but only 25% were due to topographic habitat association. Although the pronounced ridge-valley gradient and contrast of south-west versus north-east aspect created consistent habitats, our results suggest that local species assemblages at Sinharaja forest are jointly shaped by neutral and niche processes.
Understanding how plant species coexist in tropical rainforests is one of the biggest challenges in community ecology. One prominent hypothesis suggests that rare species are at an advantage because ...trees have lower survival in areas of high conspecific density due to increased attack by natural enemies, a process known as negative density dependence (NDD). A consensus is emerging that NDD is important for plant-species coexistence in tropical forests. Most evidence comes from short-term studies, but testing the prediction that NDD decreases the spatial aggregation of tree populations provides a long-term perspective. While spatial distributions have provided only weak evidence for NDD so far, the opposing effects of environmental heterogeneity might have confounded previous analyses. Here we use a novel statistical technique to control for environmental heterogeneity while testing whether spatial aggregation decreases with tree size in four tropical forests. We provide evidence for NDD in 22% of the 139 tree species analyzed and show that environmental heterogeneity can obscure the spatial signal of NDD. Environmental heterogeneity contributed to aggregation in 84% of species. We conclude that both biotic interactions and environmental heterogeneity play crucial roles in shaping tree dynamics in tropical forests.
The assertion that the spatial location of different species is independent of each other is fundamental in major ecological theories such as neutral theory that describes a stochastic geometry of ...biodiversity. However, this assertion has rarely been tested. Here we use techniques of spatial point pattern analysis to conduct a comprehensive test of the independence assertion by analysing data from three large forest plots with different species richness: a species-rich tropical forest at Barro Colorado Island (Panama), a tropical forest in Sinharaja (Sri Lanka), and a temperate forest in Changbaishan (China). We hypothesize that stochastic dilution effects owing to increasing species richness overpower signals of species associations, thereby yielding approximate species independence. Indeed, the proportion of species pairs showing: (i) no significant interspecific association increased with species richness, (ii) segregation decreased with species richness, and (iii) small-scale interspecific interaction decreased with species richness. This suggests that independence may indeed be a good approximation in the limit of very species-rich communities. Our findings are a step towards a better understanding of factors governing species-rich communities and we propose a hypothesis to explain why species placement in species-rich communities approximates independence.
A persistent challenge in ecology is to explain the high diversity of tree species in tropical forests. Although the role of species characteristics in maintaining tree diversity in tropical forests ...has been the subject of theory and debate for decades, spatial patterns in local diversity have not been analyzed from the viewpoint of individual species. To measure scale-dependent local diversity structures around individual species, we propose individual species-area relationships (ISAR), a spatial statistic that marries common species-area relationships with Ripley's K to measure the expected α diversity in circular neighborhoods with variable radius around an arbitrary individual of a target species. We use ISAR to investigate if and at which spatial scales individual species increase in tropical forests' local diversity (accumulators), decrease local diversity (repellers), or behave neutrally. Our analyses of data from Barro Colorado Island (Panama) and Sinharaja (Sri Lanka) reveal that individual species leave identifiable signatures on spatial diversity, but only on small spatial scales. Most species showed neutral behavior outside neighborhoods of 20 m. At short scales (<20 m), we observed, depending on the forest type, two strongly different roles of species: diversity repellers dominated at Barro Colorado Island and accumulators at Sinharaja. Nevertheless, we find that the two tropical forests lacked any key species structuring species diversity at larger scales, suggesting that "balanced" species-species interactions may be a characteristic of these species-rich forests. We anticipate our analysis method will be a starting point for more powerful investigations of spatial structures in diversity to promote a better understanding of biodiversity in tropical forests.
Interactions among neighboring individuals influence plant performance and should create spatial patterns in local community structure. In order to assess the role of large trees in generating ...spatial patterns in local species richness, we used the individual species-area relationship (ISAR) to evaluate the species richness of trees of different size classes (and dead trees) in circular neighborhoods with varying radius around large trees of different focal species. To reveal signals of species interactions, we compared the ISAR function of the individuals of focal species with that of randomly selected nearby locations. We expected that large trees should strongly affect the community structure of smaller trees in their neighborhood, but that these effects should fade away with increasing size class. Unexpectedly, we found that only few focal species showed signals of species interactions with trees of the different size classes and that this was less likely for less abundant focal species. However, the few and relatively weak departures from independence were consistent with expectations of the effect of competition for space and the dispersal syndrome on spatial patterns. A noisy signal of competition for space found for large trees built up gradually with increasing life stage; it was not yet present for large saplings but detectable for intermediates. Additionally, focal species with animal-dispersed seeds showed higher species richness in their neighborhood than those with gravity- and gyration-dispersed seeds. Our analysis across the entire ontogeny from recruits to large trees supports the hypothesis that stochastic effects dilute deterministic species interactions in highly diverse communities. Stochastic dilution is a consequence of the stochastic geometry of biodiversity in species-rich communities where the identities of the nearest neighbors of a given plant are largely unpredictable. While the outcome of local species interactions is governed for each plant by deterministic fitness and niche differences, the large variability of competitors causes also a large variability in the outcomes of interactions and does not allow for strong directed responses at the species level. Collectively, our results highlight the critical effect of the stochastic geometry of biodiversity in structuring local spatial patterns of tropical forest diversity.
Both habitat filtering and dispersal limitation influence the compositional structure of forest communities, but previous studies examining the relative contributions of these processes with ...variation partitioning have primarily used topography to represent the influence of the environment. Here, we bring together data on both topography and soil resource variation within eight large (24–50 ha) tropical forest plots, and use variation partitioning to decompose community compositional variation into fractions explained by spatial, soil resource and topographic variables. Both soil resources and topography account for significant and approximately equal variation in tree community composition (9–34% and 5–29%, respectively), and all environmental variables together explain 13–39% of compositional variation within a plot. A large fraction of variation (19–37%) was spatially structured, yet unexplained by the environment, suggesting an important role for dispersal processes and unmeasured environmental variables. For the majority of sites, adding soil resource variables to topography nearly doubled the inferred role of habitat filtering, accounting for variation in compositional structure that would previously have been attributable to dispersal. Our results, illustrated using a new graphical depiction of community structure within these plots, demonstrate the importance of small-scale environmental variation in shaping local community structure in diverse tropical forests around the globe.
Niche and neutral theories emphasize different processes that contribute to the maintenance of species diversity and should leave different spatial structures in species assemblages. In this study we ...used variation partitioning in combination with distance-based Moran's eigenvector maps and habitat variables to determine the relative importance of the effects of pure habitat, pure spatial, and spatially structured habitat processes on the spatial distribution of tree species composition and richness in a 25-ha tropical rain forest of Sinharaja/Sri Lanka. We analyzed the contribution of those components at three spatial scales (10 m, 20 m, and 50 m) for all trees and the three life stages: recruits, juveniles, and adults. At the 10-m scale, 80% of the variation in species composition remained unexplained for recruits and adults, but only 55% for juveniles. With increasingly broader scales these figures were strongly reduced, mainly by an increasing contribution of the spatially structured habitat component, which explained 4-30%, 20-47%, and 8-35% of variation in species composition for recruits, juveniles, and adults, respectively. The pure spatial component was most important at the 20-m scale and reached 20%, 32%, and 23% for recruits, juveniles, and adults, respectively. The spatially structured habitat component described variability at broader scales than the pure spatial component. Our results suggest that stochastic processes and spatially structuring processes of community dynamics, such as dispersal limitation and habitat association, contributed jointly to explain species composition and richness at the Sinharaja forest, but their relative importance changed with scale and life stage. Species assembly at the local scale was more strongly impacted by stochasticity, whereas the signal of habitat was stronger at the 50-m scale where plant-scale stochasticity is averaged out. Recent research points to an emerging consensus on the relative contribution of stochasticity, habitat, and spatial processes in governing community assembly, but how these components change with life stage, and how this is influenced by sample size, remains to be explored.
Studies of forest dynamics plots (FDPs) have revealed a variety of negative density-dependent (NDD) demographic interactions, especially among conspecific trees. These interactions can affect growth ...rate, recruitment and mortality, and they play a central role in the maintenance of species diversity in these complex ecosystems. Here we use an equal area annulus (EAA) point-pattern method to comprehensively analyze data from two tropical FDPs, Barro Colorado Island in Panama and Sinharaja in Sri Lanka. We show that these NDD interactions also influence the continued evolutionary diversification of even distantly related tree species in these FDPs. We examine the details of a wide range of these interactions between individual trees and the trees that surround them. All these interactions, and their cumulative effects, are strongest among conspecific focal and surrounding tree species in both FDPs. They diminish in magnitude with increasing phylogenetic distance between heterospecific focal and surrounding trees, but do not disappear or change the pattern of their dependence on size, density, frequency or physical distance even among the most distantly related trees. The phylogenetic persistence of all these effects provides evidence that interactions between tree species that share an ecosystem may continue to promote adaptive divergence even after the species' gene pools have become separated. Adaptive divergence among taxa would operate in stark contrast to an alternative possibility that has previously been suggested, that distantly related species with dispersal-limited distributions and confronted with unpredictable neighbors will tend to converge on common strategies of resource use. In addition, we have also uncovered a positive density-dependent effect: growth rates of large trees are boosted in the presence of a smaller basal area of surrounding trees. We also show that many of the NDD interactions switch sign rapidly as focal trees grow in size, and that their cumulative effect can strongly influence the distributions and species composition of the trees that surround the focal trees during the focal trees' lifetimes.
Tropical forests are notable for their high species diversity, even on small spatial scales, and right‐skewed species and size abundance distributions. The role of individual species as drivers of ...the spatial organization of diversity in these forests has been explained by several hypotheses and processes, for example, stochastic dilution, negative density dependence, or gap dynamics. These processes leave a signature in spatial distribution of small trees, particularly in the vicinity of large trees, likely having stronger effects on their neighbors. We are exploring species diversity patterns within the framework of various diversity‐generating hypotheses using individual species–area relationships. We used the data from three tropical forest plots (Wanang—Papua New Guinea, Barro Colorado Island—Panama, and Sinharaja—Sri Lanka) and included also the saplings (DBH ≥ 1 cm). Resulting cross‐size patterns of species richness and evenness reflect the dynamics of saplings affected by the distribution of large trees. When all individuals with DBH ≥1 cm are included, ~50% of all tree species from the 25‐ or 50‐ha plot can be found within 35 m radius of an individual tree. For all trees, 72%–78% of species were identified as species richness accumulators, having more species present in their surroundings than expected by null models. This pattern was driven by small trees as the analysis of DBH >10 cm trees showed much lower proportion of accumulators, 14%–65% of species identified as richness repellers and had low richness of surrounding small trees. Only 11%–26% of species had lower species evenness than was expected by null models. High proportions of species richness accumulators were probably due to gap dynamics and support Janzen–Connell hypothesis driven by competition or top‐down control by pathogens and herbivores. Observed species diversity patterns show the importance of including small tree size classes in analyses of the spatial organization of diversity.
Common spatial patterns of trees were observed in three tropical forest plots. Small trees were associated with increased diversity, while large trees were more often associated with low diversity. Gap dynamics and negative density‐dependent processes were expected to form the diversity patterns.
Fragmented and degraded ecosystems should be restored to ensure the biological connectivity among fragmented forest landscapes. The tropical rainforests of Southwestern Sri Lanka are highly degraded ...and fragmented due to plantation agriculture and human settlements. However, the high spatial variation in environmental factors and ecosystem functions challenge the success rate of restoration interventions. The aim of this study was to assess the vegetation composition and stand structure in relation to the spatial variation in key soil physicochemical parameters in the Endane Biodiversity Corridor that links peripheral forest reserve to the Sinharaja Rainforest Complex (SRC). The site that extends over 24 ha was classified into five land-use categories (productive tea lands, marginal tea lands, scrub—abandoned three years ago, and two woodlands—abandoned 15 years ago) in which the vegetation composition, stand structure, and physicochemical parameters of soil were assessed and mapped. Results revealed that the Shannon diversity index in the scrub and the woodlands were higher than in the tea lands. The diversity among the secondary forest patches was similar. However, with a mean record of 14 species, the species richness was high in sites close to the SRC. In comparison to the SRC (358 Mg ha−1), there was a substantial potential to sequester more carbon in the restoration sites (12–108 Mg ha−1). While explaining 31% of abundance and species distribution, the ordination results revealed a close relationship of the soil parameters to vegetation composition and species abundance. The calculated coefficient variation values for soil parameters (TN, EC, Av.P, Ex.K, OC, and BD) were beyond 12%, indicating high or moderate soil spatial variability among the land use categories. Coefficient of variation for soil pH was estimated to be 9%, revealing low soil spatial variability among the land use categories. The maps of these soil parameters corresponded with the type of land use and fertilizer application to tea fields. The highest and the lowest total N contents were observed in the scrub and woodlands, respectively, which appears to be mediated by the relative composition of N-fixing trees between the two groups. Our results facilitate effective matching of sites to species for restoration of the Endane Biodiversity Corridor that may be replicated in similar restoration contexts in tropical Asia.
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