Evidence that niche specialization explains species-energy relationships in lake fish communities.

1. Interspecific niche differences have long been identified as a major explanation for the occurrence of species-rich communities. However, much fieldwork studying variation in local species richness has focused upon physical habitat attributes or regional factors, such as the size of the regional species pool. 2. We applied indices of functional diversity and niche overlap to data on the species niche to examine the importance of interspecific niche differentiation for species richness in French lake fish communities. We combined this information with environmental data to test generalizations of the physiological tolerance and niche specialization hypotheses for species-energy relationships. 3. We found evidence for a largely non-saturating relationship (relative to random expectation) between species richness and functional evenness (evenness of spacing between species in niche space), while functional richness (volume of niche space occupied) peaked at moderate levels of species richness and niche overlap showed an initial decrease followed by saturation. This suggests that increased niche specialization may have allowed species to coexist in the most species-rich communities. 4. We tested for evidence that increased temperature, local habitat area, local habitat diversity and immigration affected species richness via increased niche specialization. Temperature explained by far the largest amount of variation in species richness, functional diversity and niche overlap. These results, combined with the largely non-saturating species richness-functional evenness relationship, suggest that increased temperature may have permitted increased species richness by allowing increased niche specialization. 5. These results emphasize the importance of niche differences for species coexistence in species-rich communities, and indicate that the conservation of functional diversity may be vital for the maintenance of species diversity in biological communities. Our approach may be applied readily to many types of community, and at any scale, thus providing a flexible means of testing niche-based hypotheses for species richness gradients.

Functional characters combined with null models reveal inconsistency in mechanisms of species turnover in lacustrine fish communities.

Functional characters have the potential to act as indicators of species turnover between local communities. Null models provide a powerful statistical approach to test for patterns using functional character information. A combined null model/functional character approach provides the ability to distinguish between the effect of competition and environmental filtering on species turnover. We measured 13 functional characters relating directly to resource use for the fish species found in French lakes. We combined this functional character data with a null model approach to test whether co-occurring species overlapped more or less than expected at random for four primary niche axes. We used an environmentally constrained null model approach to determine if the same mechanisms were responsible for species turnover at different sections of the altitudinal gradient. Functional diversity indices were used to examine the variation in functional character diversity with altitude, as a test of the hypothesis that competitive intensity decreases with increasing environmental adversity. The unconstrained null model showed that environmental filtering was the dominant influence on species turnover between lakes. In the constrained null model, there was much less evidence for environmental filtering, emphasising the strong effect of altitude on turnover in functional character values between local communities. Different results were obtained for low-altitude and high-altitude lake subsets, with more evidence for the effect of environmental filtering being found in the high-altitude lakes. This demonstrates that different processes may influence species turnover throughout an environmental gradient. Functional diversity values showed a slight decrease with altitude, indicating that there was only weak evidence that competitive intensity decreased with increasing altitude. Variation resource availability and environmental stress probably cause the observed turnover in functional characters along the altitudinal gradient, though the effects of dispersal limitation and species introductions in high-altitude lakes cannot be ruled out.