Variable temperature regimes and wetland salinity reduce performance of juvenile wood frogs.

On a changing planet, amphibians must respond to weather events shifting in frequency and magnitude, and to how those temperature and precipitation changes interact with other anthropogenic disturbances that modify amphibian habitat. To understand how drastic changes in environmental conditions affect wood frog tadpoles, we tested five temperature manipulations, including Ambient (water temperatures tracking daily air temperatures), Elevated (+ 3 °C above ambient), Nightly (removal of nightly lows), Spike (+ 6 °C above ambient every third week), and Flux (alternating ambient and + 3 °C weekly) crossed with Low Salt (specific conductivity: 109-207 µS-cm) and High Salt (1900-2000 µS-cm). We replicated each of the ten resulting treatments four times. High-salinity conditions produced larger metamorphs than low-salinity conditions. Tadpole survival was reduced only by the Spike treatment (P = 0.017). Elevated temperatures did not shorten larval periods; time to metamorphosis did not differ among temperature treatments (P = 0.328). We retained 135 recently metamorphosed frogs in outdoor terrestrial enclosures for 10 months to investigate larval environment carryover effects. Juvenile frogs grew larger in low-density terrestrial enclosures than high density (P = 0.015) and frogs from Ambient Low Salt larval conditions grew and survived better than frogs from manipulated larval conditions. Frogs from High Salt larval conditions had lower survival than frogs from Low Salt conditions. Our results suggest that anthropogenic disturbances to larval environmental conditions can affect both larval and post-metamorphic individuals, with detrimental carryover effects of high-salinity larval conditions not emerging until the juvenile life stage.

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.

Hourly movement decisions indicate how a large carnivore inhabits developed landscapes.

The ecology of wildlife living in proximity to humans often differs from that in more natural places. Animals may perceive anthropogenic features and people as threats, exhibiting avoidance behavior, or may acclimate to human activities. As development expands globally, changes in the ecology of species in response to human phenomena may determine whether animals persist in these changing environments. We hypothesize that American black bears (Ursus americanus) persist within developed areas by effectively avoiding risky landscape features. We test this by quantifying changes in the movements of adult females from a population living within exurban and suburban development. We collected hourly GPS data from 23 individuals from 2012 to 2014 and used step-selection functions to estimate selection for anthropogenic features. Females were more avoidant of roads and highways when with cubs than without and were more responsive to increased traffic volume. As bears occupied greater housing densities, selection for housing increased, while avoidance of roads and responsiveness to traffic increased. Behavioral flexibility allowed bears in highly developed areas to alter selection and avoidance for anthropogenic features seasonally. These findings support the hypothesis that black bears perceive human activity as risky, and effectively avoid these risks while inhabiting developed areas. We document a high amount of individual variation in selection of anthropogenic features within the study population. Our findings suggest that initially, wildlife can successfully inhabit developed landscapes by effectively avoiding human activity. However, variation among individuals provides the capacity for population-level shifts in behavior over time.

Quantifying climate sensitivity and climate-driven change in North American amphibian communities.

Changing climate will impact species' ranges only when environmental variability directly impacts the demography of local populations. However, measurement of demographic responses to climate change has largely been limited to single species and locations. Here we show that amphibian communities are responsive to climatic variability, using >500,000 time-series observations for 81 species across 86 North American study areas. The effect of climate on local colonization and persistence probabilities varies among eco-regions and depends on local climate, species life-histories, and taxonomic classification. We found that local species richness is most sensitive to changes in water availability during breeding and changes in winter conditions. Based on the relationships we measure, recent changes in climate cannot explain why local species richness of North American amphibians has rapidly declined. However, changing climate does explain why some populations are declining faster than others. Our results provide important insights into how amphibians respond to climate and a general framework for measuring climate impacts on species richness.

Spatial genetic patterns indicate mechanism and consequences of large carnivore cohabitation within development.

Patterns of human development are shifting from concentrated housing toward sprawled housing intermixed with natural land cover, and wildlife species increasingly persist in close proximity to housing, roads, and other anthropogenic features. These associations can alter population dynamics and evolutionary trajectories. Large carnivores increasingly occupy urban peripheries, yet the ecological consequences for populations established entirely within urban and exurban landscapes are largely unknown. We applied a spatial and landscape genetics approach, using noninvasively collected genetic data, to identify differences in black bear spatial genetic patterns across a rural-to-urban gradient and quantify how development affects spatial genetic processes. We quantified differences in black bear dispersal, spatial genetic structure, and migration between differing levels of development within a population primarily occupying areas with >6 houses/km2 in western Connecticut. Increased development disrupted spatial genetic structure, and we found an association between increased housing densities and longer dispersal. We also found evidence that roads limited gene flow among bears in more rural areas, yet had no effect among bears in more developed ones. These results suggest dispersal behavior is condition-dependent and indicate the potential for landscapes intermixing development and natural land cover to facilitate shifts toward increased dispersal. These changes can affect patterns of range expansion and the phenotypic and genetic composition of surrounding populations. We found evidence that subpopulations occupying more developed landscapes may be sustained by male-biased immigration, creating potentially detrimental demographic shifts.

Range position and climate sensitivity: The structure of among-population demographic responses to climatic variation.

Species' distributions will respond to climate change based on the relationship between local demographic processes and climate and how this relationship varies based on range position. A rarely tested demographic prediction is that populations at the extremes of a species' climate envelope (e.g., populations in areas with the highest mean annual temperature) will be most sensitive to local shifts in climate (i.e., warming). We tested this prediction using a dynamic species distribution model linking demographic rates to variation in temperature and precipitation for wood frogs (Lithobates sylvaticus) in North America. Using long-term monitoring data from 746 populations in 27 study areas, we determined how climatic variation affected population growth rates and how these relationships varied with respect to long-term climate. Some models supported the predicted pattern, with negative effects of extreme summer temperatures in hotter areas and positive effects on recruitment for summer water availability in drier areas. We also found evidence of interacting temperature and precipitation influencing population size, such as extreme heat having less of a negative effect in wetter areas. Other results were contrary to predictions, such as positive effects of summer water availability in wetter parts of the range and positive responses to winter warming especially in milder areas. In general, we found wood frogs were more sensitive to changes in temperature or temperature interacting with precipitation than to changes in precipitation alone. Our results suggest that sensitivity to changes in climate cannot be predicted simply by knowing locations within the species' climate envelope. Many climate processes did not affect population growth rates in the predicted direction based on range position. Processes such as species-interactions, local adaptation, and interactions with the physical landscape likely affect the responses we observed. Our work highlights the need to measure demographic responses to changing climate.

Camera trap arrays improve detection probability of wildlife: Investigating study design considerations using an empirical dataset.

Camera trapping is a standard tool in ecological research and wildlife conservation. Study designs, particularly for small-bodied or cryptic wildlife species often attempt to boost low detection probabilities by using non-random camera placement or baited cameras, which may bias data, or incorrectly estimate detection and occupancy. We investigated the ability of non-baited, multi-camera arrays to increase detection probabilities of wildlife. Study design components were evaluated for their influence on wildlife detectability by iteratively parsing an empirical dataset (1) by different sizes of camera arrays deployed (1-10 cameras), and (2) by total season length (1-365 days). Four species from our dataset that represented a range of body sizes and differing degrees of presumed detectability based on life history traits were investigated: white-tailed deer (Odocoileus virginianus), bobcat (Lynx rufus), raccoon (Procyon lotor), and Virginia opossum (Didelphis virginiana). For all species, increasing from a single camera to a multi-camera array significantly improved detection probability across the range of season lengths and number of study sites evaluated. The use of a two camera array increased survey detection an average of 80% (range 40-128%) from the detection probability of a single camera across the four species. Species that were detected infrequently benefited most from a multiple-camera array, where the addition of up to eight cameras produced significant increases in detectability. However, for species detected at high frequencies, single cameras produced a season-long (i.e, the length of time over which cameras are deployed and actively monitored) detectability greater than 0.75. These results highlight the need for researchers to be critical about camera trap study designs based on their intended target species, as detectability for each focal species responded differently to array size and season length. We suggest that researchers a priori identify target species for which inference will be made, and then design camera trapping studies around the most difficult to detect of those species.

Quantitative evidence for the effects of multiple drivers on continental-scale amphibian declines.

Since amphibian declines were first proposed as a global phenomenon over a quarter century ago, the conservation community has made little progress in halting or reversing these trends. The early search for a "smoking gun" was replaced with the expectation that declines are caused by multiple drivers. While field observations and experiments have identified factors leading to increased local extinction risk, evidence for effects of these drivers is lacking at large spatial scales. Here, we use observations of 389 time-series of 83 species and complexes from 61 study areas across North America to test the effects of 4 of the major hypothesized drivers of declines. While we find that local amphibian populations are being lost from metapopulations at an average rate of 3.79% per year, these declines are not related to any particular threat at the continental scale; likewise the effect of each stressor is variable at regional scales. This result - that exposure to threats varies spatially, and populations vary in their response - provides little generality in the development of conservation strategies. Greater emphasis on local solutions to this globally shared phenomenon is needed.

Snow cover and late fall movement influence wood frog survival during an unusually cold winter.

Understanding how organisms will respond to altered winter conditions is hampered by a paucity of information on the winter ecology for many species. Amphibians are sensitive to environmental temperature and moisture conditions and may be vulnerable to changes in winter climate. We used a combination of radio telemetry and field enclosures to monitor survival of the freeze-tolerant wood frog (Lithobates sylvaticus) during the unusually cold winter of 2013-2014. We experimentally manipulated snow cover to determine the effect of snow removal on winter survival. In addition, we placed a group of untracked frogs at locations used by tracked frogs prior to long-distance late fall movement to investigate whether late fall movement entailed survival consequences. Winter survival was highest (75.3 %) among frogs at post-movement locations that received natural snow cover. The odds of surviving the winter for frogs in the snow removal treatment was only 21.6 % that of frogs in the natural snow treatment. Likewise, paired frogs placed at pre-fall movement locations had only 35.1 % the odds of surviving as tracked frogs at post-fall movement locations. A comparison of a priori models that included microhabitat conditions measured at wood frog overwintering locations revealed that the minimum temperature experienced and the depth of the frog in the substrate explained additional variation in winter survival. Our results suggest that acute exposure to lethal temperature conditions is the most likely cause of mortality during this study, rather than energy exhaustion or desiccation. They also demonstrate the importance of snow cover to the winter survival of wood frogs.

Demographics and density estimates of two three-toed box turtle (Terrapene carolina triunguis) populations within forest and restored prairie sites in central Missouri.

Box turtles (Terrapene carolina) are widely distributed but vulnerable to population decline across their range. Using distance sampling, morphometric data, and an index of carapace damage, we surveyed three-toed box turtles (Terrapene carolina triunguis) at 2 sites in central Missouri, and compared differences in detection probabilities when transects were walked by one or two observers. Our estimated turtle densities within forested cover was less at the Thomas S. Baskett Wildlife Research and Education Center, a site dominated by eastern hardwood forest (d = 1.85 turtles/ha, 95% CI [1.13, 3.03]) than at the Prairie Fork Conservation Area, a site containing a mix of open field and hardwood forest (d = 4.14 turtles/ha, 95% CI [1.99, 8.62]). Turtles at Baskett were significantly older and larger than turtles at Prairie Fork. Damage to the carapace did not differ significantly between the 2 populations despite the more prevalent habitat management including mowing and prescribed fire at Prairie Fork. We achieved improved estimates of density using two rather than one observer at Prairie Fork, but negligible differences in density estimates between the two methods at Baskett. Error associated with probability of detection decreased at both sites with the addition of a second observer. We provide demographic data on three-toed box turtles that suggest the use of a range of habitat conditions by three-toed box turtles. This case study suggests that habitat management practices and their impacts on habitat composition may be a cause of the differences observed in our focal populations of turtles.

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.

Survival costs associated with wood frog breeding migrations: effects of timber harvest and drought.

Migration presents a trade-off for individuals between the potential fitness benefits of reaching high-quality habitat vs. the potential costs of migration. Within an information-theoretic framework, we examined the costs of migration for adult wood frogs (Rana sylvatica) in response to timber harvest and annual weather conditions using Cox proportional-hazard estimates of survival. In 2004 prior to timber harvest, survival did not differ between the inside (0.75, SE = 0.078) and outside (0.73, SE = 0.235) of the circular timber harvest arrays (each 164 m radius). Following timber harvest, survival inside arrays in both 2005 and 2006 (0.22, SE = 0.065; 0.42, SE = 0.139) was lower than survival outside of the arrays and prior to harvest. Sources of mortality included predation in all years and desiccation in the drought year of 2005. The most-supported models for explaining both predation and desiccation risks reflected behaviors as opposed to timber harvest or weather conditions. Both predation and desiccation risks increased when frogs made frequent movements or were located near breeding ponds. Optimal behaviors for reducing predation and desiccation risks were the same before and after timber harvest; however, the survival consequences for not adopting these behaviors were more severe following harvest. Our results provide empirical evidence for (1) the ecological pressures that influence migratory behavior and (2) differential survival in relation to migratory behavior which reveals why frogs move relatively long distances away from breeding sites.

Demographic consequences of terrestrial habitat loss for pool-breeding amphibians: predicting extinction risks associated with inadequate size of buffer zones.

Much of the biodiversity associated with isolated wetlands requires aquatic and terrestrial habitat to maintain viable populations. Current federal wetland regulations in the United States do not protect isolated wetlands or extend protection to surrounding terrestrial habitat. Consequently, some land managers, city planners, and policy makers at the state and local levels are making an effort to protect these wetland and neighboring upland habitats. Balancing human land-use and habitat conservation is challenging, and well-informed land-use policy is hindered by a lack of knowledge of the specific risks of varying amounts of habitat loss. Using projections of wood frog (Rana sylvatica) and spotted salamander (Ambystoma maculatum) populations, we related the amount of high-quality terrestrial habitat surrounding isolated wetlands to the decline and risk of extinction of local amphibian populations. These simulations showed that current state-level wetland regulations protecting 30 m or less of surrounding terrestrial habitat are inadequate to support viable populations of pool-breeding amphibians. We also found that species with different life-history strategies responded differently to the loss and degradation of terrestrial habitat. The wood frog, with a short life span and high fecundity, was most sensitive to habitat loss and isolation, whereas the longer-lived spotted salamander with lower fecundity was most sensitive to habitat degradation that lowered adult survival rates. Our model results demonstrate that a high probability of local amphibian population persistence requires sufficient terrestrial habitat, the maintenance of habitat quality, and connectivity among local populations. Our results emphasize the essential role of adequate terrestrial habitat to the maintenance of wetland biodiversity and ecosystem function and offer a means of quantifying the risks associated with terrestrial habitat loss and degradation.

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.

Leaf litter input mediates tadpole performance across forest canopy treatments.

Understanding the mechanisms limiting the distributions of organisms is necessary for predicting changes in community composition along habitat gradients. In many areas of the USA, land originally cleared for agriculture has been undergoing a process of reforestation, creating a gradient of canopy cover. For small temporary wetlands, this gradient can alter abiotic conditions and influence the resource base of wetland food webs by affecting litter inputs. As distributions of amphibians and many other temporary wetland taxa correlate with canopy cover, we experimentally manipulated shade levels and litter types in pond mesocosms to explore mechanisms limiting species performance in wetlands with canopy cover. Most differences between ponds were mediated by litter type rather than direct effects of shading. Although all three amphibian species tested are open-canopy specialists, spring peepers were the only species to show decreased survival in shaded ponds. Pond litter type generally had strong effects on growth and development rates, with tadpoles of two species in grass litter ponds growing to twice the size of, and metamorphosing 7 days earlier than, those in leaf litter ponds. Contrary to our initial hypothesis, shade level and litter type showed very few significant interactions. Our results indicate that the effects of shading cannot be considered in isolation of vegetation changes in pond basins when evaluating the effects of forest succession on temporary pond communities.