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

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.

Post-Pleistocene differentiation in a Central Interior Highlands endemic salamander.

AIM: For many endemic species with limited dispersal capacities, the relationship between landscape changes and species distributions is still unclear. We characterized the population structure of the endemic ringed salamander (Ambystoma annulatum) across its distribution in the Central Interior Highlands (CIH) of North America, an area of high species endemism, to infer the ecological and evolutionary history of the species. METHODS: We sampled 498 individuals across the species distribution and characterized the population genetic structure using nuclear microsatellite and mitochondrial DNA (mtDNA) markers. RESULTS: Ambystoma annulatum exist in two strongly supported nuclear genetic clusters across the CIH that correspond to a northern cluster that includes the Missouri Ozark populations and a southern cluster that includes the Arkansas and Oklahoma Ozarks and the Ouachita Mountains. Our demographic models estimated that these populations diverged approximately 2,700 years ago. Pairwise estimates of genetic differentiation at microsatellite and mtDNA markers indicated limited contemporary gene flow and suggest that genetic differentiation was primarily influenced by changes in the post-Pleistocene landscape of the CIH. MAIN CONCLUSIONS: Both the geologic history and post-European settlement history of the CIH have influenced the population genetic structure of A. annulatum. The low mtDNA diversity suggests a retraction into and expansion out of refugial areas in the south-central Ozarks, during temperature fluctuations of the Pleistocene and Holocene epochs. Similarly, the estimated divergence time for the two nuclear clusters corresponds to changes in the post-Pleistocene landscape. More recently, decreased A. annulatum gene flow may be a result of increased habitat fragmentation and alteration post-European settlement.

The influence of breeding phenology on the genetic structure of four pond-breeding salamanders.

Understanding metapopulation dynamics requires knowledge about local population dynamics and movement in both space and time. Most genetic metapopulation studies use one or two study species across the same landscape to infer population dynamics; however, using multiple co-occurring species allows for testing of hypotheses related to different life history strategies. We used genetic data to study dispersal, as measured by gene flow, in three ambystomatid salamanders (Ambystoma annulatum, A. maculatum, and A. opacum) and the Central Newt (Notophthalmus viridescens louisianensis) on the same landscape in Missouri, USA. While all four salamander species are forest dependent organisms that require fishless ponds to reproduce, they differ in breeding phenology and spatial distribution on the landscape. We use these differences in life history and distribution to address the following questions: (1) Are there species-level differences in the observed patterns of genetic diversity and genetic structure? and (2) Is dispersal influenced by landscape resistance? We detected two genetic clusters in A. annulatum and A. opacum on our landscape; both species breed in the fall and larvae overwinter in ponds. In contrast, no structure was evident in A. maculatum and N. v. louisianensis, species that breed during the spring. Tests for isolation by distance were significant for the three ambystomatids but not for N. v. louisianensis. Landscape resistance also contributed to genetic differentiation for all four species. Our results suggest species-level differences in dispersal ability and breeding phenology are driving observed patterns of genetic differentiation. From an evolutionary standpoint, the observed differences in dispersal distances and genetic structure between fall breeding and spring breeding species may be a result of the trade-off between larval period length and size at metamorphosis which in turn may influence the long-term viability of the metapopulation. Thus, it is important to consider life history differences among closely related and ecologically similar species when making management decisions.