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Amphibian species richness across environmental gradients


  • Earl E. Werner,

  • David K. Skelly,

  • Rick A. Relyea,

  • Kerry L. Yurewicz

E. E. Werner (, Dept of Ecology and Evolutionary Biology, Univ. of Michigan, Ann Arbor, MI 48109, USA. – D. K. Skelly, School of Forestry & Environmental Studies, Yale Univ., New Haven, CT 06511, USA. – R. A. Relyea, Dept of Biological Sciences, Univ. of Pittsburgh, Pittsburgh, PA 15260, USDA. – K. L. Yurewicz, Biological Sciences, Plymouth State Univ., Plymouth, NH 03264, USA.


Large-scale field patterns are a fundamental source of inferences on processes responsible for variation in species richness among habitats. We examined species richness of larval amphibian communities in 37 ponds over seven years on the Univ. of Michigan's E. S. George Reserve. Ordination of the community incidence matrix indicated a strong major axis of variation in species associations that was correlated with pond hydroperiod, surface area and forest canopy cover. Communities were significantly nested with those species found in ponds with high canopy cover, small area and short hydroperiod being nested subsets of those found in ponds with contrasting characteristics. Presence of fish had strong negative effects on species richness; relaxation of this effect also was apparent when fish were extirpated from ponds by drought. We employed a model selection analysis to identify the most appropriate statistical model for predicting the long-term average species richness of these ponds from local abiotic and biotic (predator and competitor density) factors. A model including only the abiotic factors was overwhelmingly superior for the anurans; hierarchical partitioning indicated that area and canopy cover alone accounted for over 70% of the independent effects of predictor variables. The global model including both abiotic and biotic factors was the best supported model for the caudates, and correspondingly hierarchical partitioning suggested that area, hydroperiod, invertebrate predators and caudate biomass all accounted for 9–16% of the independent effects. Overall, biotic factors accounted for much less of the variation in species richness than abiotic factors. The patterns in larger, open-canopy ponds provided little evidence of competitive effects on species richness, though there were patterns consistent with competitive effects in small, closed-canopy ponds. The unusual temporal and spatial extent of these data enabled us to critically evaluate ideas regarding patterns in larval amphibian communities, and the effects of area, disturbance (hydroperiod) and productivity (canopy cover) on species richness of these communities. These results have important implications to the conservation of amphibian species richness in freshwater wetlands, which are among the most threatened ecosystems worldwide.