A central current debate in community ecology concerns the relative importance of deterministic versus stochastic processes underlying community structure. However, the concept of stochasticity presents several profound philosophical, theoretical and empirical challenges, which we address here. The philosophical argument that nothing in nature is truly stochastic can be met with the following operational concept of neutral stochasticity in community ecology: change in the composition of a community (i.e. community dynamics) is neutrally stochastic to the degree that individual demographic events -birth, death, immigration, emigration -which cause such changes occur at random with respect to species identities. Empirical methods for identifying the stochastic component of community dynamics or structure include null models and multivariate statistics on observational species-by-site data (with or without environmental or trait data), and experimental manipulations of 'stochastic' species colonization order or relative densities and frequencies of competing species. We identify the fundamental limitations of each method with respect to its ability to allow inferences about stochastic community processes. Critical future needs include greater precision in articulating the link between results and ecological inferences, a comprehensive theoretical assessment of the interpretation of statistical analyses of observational data, and experiments focusing on community size and on natural variation in species colonization order.
Abstract. Tropical montane species are characterized by narrow elevational distributions. Recent perspectives on mechanisms maintaining these restricted distributions have emphasized abiotic processes, but biotic processes may also play a role in their establishment or maintenance. One historically popular hypothesis, especially for birds, is that interspecific competition constrains ranges of closely related species that ''replace'' each other along elevational gradients. Supporting evidence, however, is based on patterns of occurrence and does not reveal potential mechanisms. We experimentally tested a prediction of this hypothesis in two genera of tropical songbirds, Catharus (Turdidae) and Henicorhina (Troglodytidae), in which species have nonoverlapping elevational distributions. Using heterospecific playback trials, we found that individuals at replacement zones showed aggressive territorial behavior in response to songs of congeners. As distance from replacement zones increased, aggression toward congener song decreased, suggesting a learned component to interspecific aggression. Additionally, aggressive responses in Catharus were asymmetric, indicating interspecific dominance. These results provide experimental evidence consistent with the hypothesis that interspecific competitive interactions restrict ranges of Neotropical birds. Our results also underscore the need to consider biotic processes, such as competition, when predicting how species' ranges will shift with climate change. Asymmetric aggression could be particularly important. For example, if warming in montane landscapes allows upslope range expansion by dominant competitors, then high-elevation subordinate species could be forced into progressively smaller mountaintop habitats, jeopardizing viability of their populations.
Summary 1.Understanding how species in a diverse regional pool are spatially distributed with respect to habitat types is a longstanding problem in ecology. Tropical species are expected to be specialists along environmental gradients, and this should result in rapid compositional change (high beta diversity) across landscapes, particularly when alpha diversity is a small fraction of regional diversity. Corollary challenges are then to identify controlling environmental variables and to ask whether species cluster into discrete community types along a gradient. 2. We investigated patterns of avian species' distributions in the Tilarán mountains of Costa Rica between 1000 m and 1700 m elevation where a strong moisture gradient exists. High beta diversity was found with both auditory counts adjusted for detectability and extensive capture data, revealing nearly complete change in community composition over a few kilometres on the Pacific slope. As predicted, this beta diversity was roughly twice as high as on temperate mountainsides. 3. Partial Mantel analyses and canonical correspondence analysis indicate that change in species composition is highly correlated with change in moisture (and correlated epiphyte cover) at different distances from the continental divide on the Pacific slope. Altitude was not a good predictor of change in species composition, as species composition varies substantially among sites at the same elevation. 4. Detrended correspondence analysis and cluster analysis revealed a zone of rapid transition separating a distinct cloud forest community from rainshadow forest. On the Caribbean slope, where a shallower moisture gradient was predicted to result in lower beta diversity, we found lower rates of compositional change and more continuous species turnover. 5. Results suggest that habitat specialization of birds is likely a strong ecological force generating high beta diversity in montane landscapes. Despite overall rapid rates of species turnover, zones of relatively coherent composition could be identified. 6. Landscapes with such high beta diversity are common in the tropics, although little studied. They offer high benefit/cost opportunities for conservation, particularly as climate change threatens to alter the species composition of communities of habitat specialists.
Aim Understanding patterns of species turnover along environmental gradients and their consistency across taxonomic groups is central to the study of biodiversity. We may expect congruence in diversity patterns across groups whose ranges could be influenced by species interactions. We explore associations between bird and vegetation communities in the tropical Andes to determine whether patterns of species richness and turnover in birds and trees are congruent with elevation, and whether tree species composition, vegetation structure, elevation, or a combination of these best predicts bird species composition. Location A forested 2600‐m elevational gradient on the eastern slope of the Peruvian Andes. Methods Bird surveys and vegetation measurements were conducted at 172 points, and a subset of these were spatially matched with fourteen 1‐ha tree inventory plots. Diversity patterns were described for trees, birds, and avian foraging guilds. We used dissimilarity matrices to examine patterns of turnover and nestedness. Turnover of birds and trees was examined by comparing compositional change of adjacent plots along the gradient. Multiple regression on distance matrices was employed to determine contributions of tree species composition, vegetation structure and elevation to explaining variation in bird species composition. Results Species richness was higher for trees than for birds, and whereas diversity in both taxa decreased with elevation, tree richness showed a low‐elevation plateau before declining at higher elevations. Tree species had narrower distributions compared to birds, but patterns of turnover were largely congruent between taxa. Nestedness contributed much less to dissimilarity than turnover, although birds showed higher nestedness, particularly at high elevations. Tree species composition, elevation and vegetation structure were all important predictors of bird species composition; the best model explained 78% of bird dissimilarity across plots. Tree species composition was always included in the best models, for all birds and foraging guilds. Main conclusions Our assessment of Andean bird and vegetation communities suggests strong correspondence, perhaps due to direct interactions or similar underlying drivers. We hypothesize that with climate change, range shifts in these groups may not occur independently. Rather, birds may have delayed upslope shifts or may be limited to high‐elevation patches where appropriate vegetation communities exist.
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