Demographic effects of phenological variation in natural populations of two pond-breeding salamanders.

Phenology is a key driver of population and community dynamics. Phenological metrics (e.g., first date that an event occurred) often simplify information from the full phenological distribution, which may undermine efforts to determine the importance of life history events. Data regarding full phenological distributions are especially needed as many species are shifting phenology with climatic change which can alter life-history patterns and species dynamics. We tested whether skewness, kurtosis or maximum duration of breeding phenology affected juvenile emigration phenology and survival in natural populations of ringed (Ambystoma annulatum) and spotted salamanders (A. maculatum) spanning a 7-year period at two study locations. We evaluated the relative importance of different phenological metrics in breeding phenology and larval density dependence on emigration phenology and survival. We found that variability in emigration phenology differed by species, with ringed salamanders having a shorter duration and distributions that were more often right-skewed and leptokurtic compared to spotted salamanders. Emigration phenology was not linked to any measure of variability in breeding phenology, indicating phenological variability operates independently across life stages and may be subject to stage-specific influences. Emigration duration and skewness were partially explained by larval density, which demonstrates how phenological distributions may change with species interactions. Further tests that use the full phenological distribution to link variability in timing of life history events to demographic traits such as survival are needed to determine if and how phenological shifts will impact species persistence.

Abundance and phenology patterns of two pond-breeding salamanders determine species interactions in natural populations.

Phenology often determines the outcome of interspecific interactions, where early-arriving species often dominate interactions over those arriving later. The effects of phenology on species interactions are especially pronounced in aquatic systems, but the evidence is largely derived from experimental studies. We examined whether differences in breeding phenology between two pond-breeding salamanders (Ambystoma annulatum and A. maculatum) affected metamorph recruitment and demographic traits within natural populations, with the expectation that the fall-breeding A. annulatum would negatively affect the spring-breeding A. maculatum. We monitored populations of each species at five ponds over 4 years using drift fences. Metamorph abundance and survival of A. annulatum were affected by intra- and interspecific processes, whereas metamorph size and date of emigration were primarily influenced by intraspecific effects. Metamorph abundance, snout-vent length, date of emigration and survival for A. maculatum were all predicted by combinations of intra- and interspecific effects, but often showed negative relationships with A. annulatum metamorph traits and abundance. Size and date of metamorphosis were strongly correlated within each species, but in opposite patterns (negative for A. annulatum and positive for A. maculatum), suggesting that the two species use alternative strategies to enhance terrestrial survival and that these factors may influence their interactions. Our results match predictions from experimental studies that suggest recruitment is influenced by intra- and interspecific processes which are determined by phenological differences between species. Incorporating spatiotemporal variability when modeling population dynamics is necessary to understand the importance of phenology in species interactions, especially as shifts in phenology occur under climate change.

Effects of fine-scale forest habitat quality on movement and settling decisions in juvenile pond-breeding salamanders.

A better understanding of how individuals respond to variation in habitat quality while moving through heterogeneous habitats is needed to predict ecological phenomena at larger scales, such as local population and metapopulation dynamics. We sought to identify how fine-scale habitat quality affects the decisions of juvenile pond-breeding salamanders (Ambystoma maculatum and A. annulatum) to cease dispersive movements away from their natal pond, select a refuge, and settle. Because of the acute susceptibility of juvenile amphibians to evaporative water loss in terrestrial habitats, we predicted that they possess mechanisms for adjusting their behavior in response to variations in fine-scale habitat quality. We used experimental field enclosures to isolate the effects of habitat quality on settling behavior and employed generalized linear mixed models to examine how manipulations in canopy cover (closed or open) and microhabitat (control, compacted soils, high coarse woody debris, high burrow density), along with environmental variables (rainfall and air temperature), affect the individual's probability of settling. Our results indicated that A. maculatum and A. annulatum had a 10% and 30% decreased probability of settling in open-canopy clearcut habitat, respectively, compared to closed-canopy forest habitat. In addition, A. annulatum were 24% less likely to settle in compacted soil treatments. Although the settlement probability of A. annulatum did not depend on refuge availability, A. maculatum were 18% and 25% more likely to settle under conditions of high burrow density and high coarse woody debris, respectively. These findings make a unique contribution to our understanding of amphibian movement ecology by demonstrating how the interplay of external factors and individual behavior produce observed patterns of movement and habitat selection.