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 of montane salamanders over an elevational gradient.

Climate change is expected to systematically alter the distribution and population dynamics of species around the world. The effects are expected to be particularly strong at high latitudes and elevations, and for ectothermic species with small ranges and limited movement potential, such as salamanders in the southern Appalachian Mountains. In this study, we sought to establish baseline abundance estimates for plethodontid salamanders (family: Plethodontidae) over an elevational gradient in Great Smoky Mountains National Park. In addition to generating these baseline data for multiple species, we describe methods for surveying salamanders that allow for meaningful comparisons over time by separating observation and ecological processes generating the data. We found that Plethodon jordani had a mid-elevation peak (1,500 m) in abundance and Desmognathus wrighti increased in abundance with elevation up to the highest areas of the park (2025 m), whereas Eurycea wilderae increased in abundance up to 1,600 m and then plateaued with increasing uncertainty. Litter depth, herbaceous ground cover, and proximity to stream were also important predictors of abundance (dependent upon species), whereas daily temperature, precipitation, ground cover, and humidity influenced detection rates. Our data provide some of the first minimally biased information for future studies to assess changes in the abundance and distribution of salamanders in this region. Understanding abundance patterns along with detailed baseline distributions will be critical for comparisons with future surveys to understand the population and community-level effects of climate change on montane salamanders.

Predicted alteration of surface activity as a consequence of climate change.

Wildlife are faced with numerous threats to survival, none more pressing than that of climate change. Understanding how species will respond behaviorally, physiologically, and demographically to a changing climate is a cornerstone of many contemporary ecological studies, especially for organisms, such as amphibians, whose persistence is closely tied to abiotic conditions. Activity is a useful parameter for understanding the effects of climate change because activity is directly linked to fitness as it dictates foraging times, energy budgets, and mating opportunities. However, activity can be challenging to measure directly, especially for secretive organisms like plethodontid salamanders, which only become surface active when conditions are cool and moist because of their anatomical and physiological restrictions. We estimated abiotic predictors of surface activity for the seven species of the Plethodon jordani complex. Five independent data sets collected from 2004 to 2017 were used to determine the parameters driving salamander surface activity in the present day, which were then used to predict potential activity changes over the next 80 yrs. Average active seasonal temperature and vapor pressure deficit were the strongest predictors of salamander surface activity and, without physiological or behavioral modifications, salamanders were predicted to exhibit a higher probability of surface activity during peak active season under future climate conditions. Temperatures during the active season likely do not exceed salamander thermal maxima to cause activity suppression and, until physiological limits are reached, future conditions may continue to increase activity. Our model is the first comprehensive field-based study to assess current and future surface activity probability. Our study provides insights into how a key behavior driving fitness may be affected by climate change.

A geostatistical state-space model of animal densities for stream networks.

Population dynamics are often correlated in space and time due to correlations in environmental drivers as well as synchrony induced by individual dispersal. Many statistical analyses of populations ignore potential autocorrelations and assume that survey methods (distance and time between samples) eliminate these correlations, allowing samples to be treated independently. If these assumptions are incorrect, results and therefore inference may be biased and uncertainty underestimated. We developed a novel statistical method to account for spatiotemporal correlations within dendritic stream networks, while accounting for imperfect detection in the surveys. Through simulations, we found this model decreased predictive error relative to standard statistical methods when data were spatially correlated based on stream distance and performed similarly when data were not correlated. We found that increasing the number of years surveyed substantially improved the model accuracy when estimating spatial and temporal correlation coefficients, especially from 10 to 15 yr. Increasing the number of survey sites within the network improved the performance of the nonspatial model but only marginally improved the density estimates in the spatiotemporal model. We applied this model to brook trout data from the West Susquehanna Watershed in Pennsylvania collected over 34 yr from 1981 to 2014. We found the model including temporal and spatiotemporal autocorrelation best described young of the year (YOY) and adult density patterns. YOY densities were positively related to forest cover and negatively related to spring temperatures with low temporal autocorrelation and moderately high spatiotemporal correlation. Adult densities were less strongly affected by climatic conditions and less temporally variable than YOY but with similar spatiotemporal correlation and higher temporal autocorrelation.

A hierarchical model of daily stream temperature using air-water temperature synchronization, autocorrelation, and time lags.

Water temperature is a primary driver of stream ecosystems and commonly forms the basis of stream classifications. Robust models of stream temperature are critical as the climate changes, but estimating daily stream temperature poses several important challenges. We developed a statistical model that accounts for many challenges that can make stream temperature estimation difficult. Our model identifies the yearly period when air and water temperature are synchronized, accommodates hysteresis, incorporates time lags, deals with missing data and autocorrelation and can include external drivers. In a small stream network, the model performed well (RMSE = 0.59°C), identified a clear warming trend (0.63 °C decade(-1)) and a widening of the synchronized period (29 d decade(-1)). We also carefully evaluated how missing data influenced predictions. Missing data within a year had a small effect on performance (∼0.05% average drop in RMSE with 10% fewer days with data). Missing all data for a year decreased performance (∼0.6 °C jump in RMSE), but this decrease was moderated when data were available from other streams in the network.

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 red-backed salamanders on ecosystem functions.

Ecosystems provide a vast array of services for human societies, but understanding how various organisms contribute to the functions that maintain these services remains an important ecological challenge. Predators can affect ecosystem functions through a combination of top-down trophic cascades and bottom-up effects on nutrient dynamics. As the most abundant vertebrate predator in many eastern US forests, woodland salamanders (Plethodon spp.) likely affect ecosystems functions. We examined the effects of red-backed salamanders (Plethodon cinereus) on a variety of forest ecosystem functions using a combined approach of large-scale salamander removals (314-m(2) plots) and small-scale enclosures (2 m(2)) where we explicitly manipulated salamander density (0, 0.5, 1, 2, 4 m(-2)). In these experiments, we measured the rates of litter and wood decomposition, potential nitrogen mineralization and nitrification rates, acorn germination, and foliar insect damage on red oak seedlings. Across both experimental venues, we found no significant effect of red-backed salamanders on any of the ecosystem functions. We also found no effect of salamanders on intraguild predator abundance (carabid beetles, centipedes, spiders). Our study adds to the already conflicting evidence on effects of red-backed salamander and other amphibians on terrestrial ecosystem functions. It appears likely that the impact of terrestrial amphibians on ecosystem functions is context dependent. Future research would benefit from explicitly examining terrestrial amphibian effects on ecosystem functions under a variety of environmental conditions and in different forest types.

Effects of timber harvesting on pond-breeding amphibian persistence: testing the evacuation hypothesis.

Numerous studies have documented the decline of amphibians following timber harvest. However, direct evidence concerning the mechanisms of population decline is lacking and hinders attempts to develop conservation or recovery plans and solutions for forest species. We summarized the mechanisms by which abundance of amphibians may initially decline following timber harvest into three testable hypotheses: (1) mortality, (2) retreat, and (3) evacuation. Here, we tested the evacuation hypothesis within a large-scale, replicated experiment. We used drift fences with pitfall traps to capture pond-breeding amphibians moving out of experimental clearcut quadrants and into control quadrants at four replicate arrays located within the Daniel Boone Conservation Area on the upper Ozark Plateau in Warren County, Missouri, USA. During the preharvest year of 2004, only 51.6% of the 312 individuals captured were moving out of pre-clearcut quadrants, and movement did not differ from random. In contrast, during both postharvest years of 2005 and 2006, the number of captures along the quadrant edge increased, and a higher proportion of individuals (59.9% and 56.6%, respectively, by year) were moving out of clearcut quadrants than entering. Salamanders moved out of clearcuts in large percentages (Ambystoma annulatum, 78.2% in 2005, 78.2% in 2006; A. maculatum, 64.0% in 2005, 57.1% in 2006). Frogs and toads also moved out of clearcut quadrants, but in lower percentages (Bufo americanus, 59.6% in 2005, 53.3% in 2006; Rana clamitans, 52.7% in 2006). Salamanders moved out of clearcuts with low-wood treatments more than out of clearcuts with high-wood treatments. Movement of salamanders out of clearcuts was independent of sex. Estimated movement out of clearcuts represented between 8.7% and 35.0% of the total breeding adults captured for two species of salamanders. Although we recognize that some portion of the amphibian population may retreat underground for short periods and others may not survive the effects of timber harvest, these data are the first direct evidence showing that individuals are capable of leaving clearcuts and shifting habitat use.