Direct effects influence larval salamander size and density more than indirect effects.

Abstract

Direct and indirect effects both influence population and community dynamics. The relative strengths of these pathways are often compared using experimental approaches, but their evaluation in situ has been less frequent. We examined how individual and aggregate impacts of direct and indirect effects of species densities, proxies for competition and predation pressure, and habitat variables influenced patterns of larval density and body size of ringed (Ambystoma annulatum) and spotted salamanders (A. maculatum). We surveyed > 150 ponds in Missouri, USA, from 2012 to 2014 to measure the density and body size of each focal species, the density of co-occurring pond food web members, and select habitat features. We used structural equation modeling to quantify the relative importance of direct and indirect pathways on both body size and larval density. Overall, both responses were explained through a combination of direct and indirect effects. However, the magnitudes of direct effects were often greater than indirect effects. Some of the direct and indirect relationships with larval salamander size and density were also consistent with results from experimental studies. Finally, total direct and indirect effects were often weaker due to habitat and density variables negating each other's impacts. Overall, our study shows that direct effects were equivalent to, or more important than, indirect effects. We also demonstrate that the effects stemming from individual relationships can sum to produce net patterns that are negligible in magnitude. Further work on direct and indirect effects with observational data are needed to examine their magnitudes in natural communities.