Spatial Heterogeneity of Rana boylii Habitat: Quantification and Ecological Meaningfulness

Analysis of the heterogeneity of stream habitat and how biological communities respond to that complexity are fundamental components of ecosystem analysis that are often inadequately addressed in watershed assessments and restoration practices. Many aquatic species, such as the Foothill Yellow-legged Frog (Rana boylii), known to associate with certain physical habitats at various times throughout their lifecycle may require some degree of habitat complexity at a larger reach scale for a population to persist. Recent research in the field of landscape ecology has expanded the use of spatial heterogeneity indices to other fields of ecology as an objective method to quantify variability in habitat. Provided that indices are used in an appropriate context and are shown to be ecologically meaningful, they provide a potentially useful tool for quantifying the variability in riverine habitat for aquatic species such as R. boylii. This study evaluated whether stream reaches with a high heterogeneity of geomorphic features, as measured by several key spatial heterogeneity indices, correlated with a greater relative abundance of R. boylii. R. boylii habitat associations were quantified throughout a single season to obtain further insight into the local hydraulic and geomorphic conditions preferred by each lifestage. The two best predictors of habitat associations by lifestage were velocity and substrate size, two key characteristics of geomorphic units such as riffles and pools. The heterogeneity of geomorphic units was then quantified and measured at the reach scale using a variety of spatial indices. Indices of spatial composition, such as Shannon's Diversity Index, were found to correlate well with frog abundance, while indices of spatial configuration, such as Contagion, were not significant. These findings indicate R. boylii may select stream reaches with increased geomorphic complexity that potentially provide habitats suitable to each lifestage with multiple functions and a greater variety of refugia as flows fluctuate throughout the season. The findings also show that spatial indices are useful in quantifying aquatic habitat heterogeneity, and when shown to be ecologically meaningful would be useful as an objective means to quantify habitat in river restoration practices.