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Ecosystem-based management aims to preserve old-growth forest attributes using techniques mimicking natural disturbances. One such technique is irregular shelterwood logging, but its impacts on forest floor organisms are poorly known. Our objective was to quantify the effects of three different treatments of irregular shelterwood on population structure and the body condition of the eastern red-backed salamander (Plethodon cinereus) 5–6 years following harvesting. A total of 64 sites in western Québec were sampled using artificial refugia and quadrat searches. Large salamanders (>32 mm) were more abundant in the gap treatment than in strip, uniform, or control treatments. Small salamanders (≤32 mm) followed the same pattern, although the differences were marginal. For a given treatment, small salamanders were as abundant as large salamanders. Salamander body condition differed between the 2 years of sampling but did not differ among treatments, regardless of salamander size. We conclude that environmental conditions in irregular shelterwood treatments 5–6 years following harvesting support populations of small vertebrates on the forest floor.

Climate change poses several challenges to biological communities including changes in the frequency of encounters between closely related congeners as a result of range shifts. When climate change leads to increased hybridization, hybrid dysfunction or genetic swamping may increase extinction risk—particularly in range‐restricted species with low vagility. The Peaks of Otter Salamander, Plethodon hubrichti, is a fully terrestrial woodland salamander that is restricted to ~18 km of ridgeline in the mountains of southwestern Virginia, and its range is surrounded by the abundant and widespread Eastern Red‐backed Salamander, Plethodon cinereus. In order to determine whether these two species are hybridizing and how their range limits may be shifting, we assessed variation at eight microsatellite loci and a 1,008 bp region of Cytochrome B in both species at allopatric reference sites and within a contact zone. Our results show that hybridization between P. hubrichti and P. cinereus either does not occur or is very rare. However, we find that diversity and differentiation are substantially higher in the mountaintop endemic P. hubrichti than in the widespread P. cinereus, despite similar movement ability for the two species as assessed by a homing experiment. Furthermore, estimation of divergence times between reference and contact zone populations via approximate Bayesian computation is consistent with the idea that P. cinereus has expanded into the range of P. hubrichti. Given the apparent recent colonization of the contact zone by P. cinereus, future monitoring of P. cinereus range limits should be a priority for the management of P. hubrichti populations., This study uses population genetic approaches to assess whether hybridization between a severely range‐restricted mountaintop endemic salamander and a closely related widespread congener is contributing to the extinction risk of the former. While we find little to no evidence of hybridization, we do present evidence that the more common species is expanding into the range of the mountaintop endemic species. We discuss the conservation implications of this finding.

For many amphibian species, reduced landscape connectivity results in reduced genetic connectivity among populations. However, large effective population sizes (Ne) slow the rate of genetic drift, causing subdivided populations to remain genetically similar despite little gene flow among them. Therefore, it is important to address the combined effects of Ne and matrix permeability to quantify the relative importance of gene flow and genetic drift on isolated amphibian populations. We applied a landscape genetic approach to investigate how patterns of gene flow (m), Ne (inferred via θ) and genetic differentiation differ among Eastern Red-backed Salamander (Plethodon cinereus) populations in a fragmented landscape (n = 4) compared to a continuous forest (n = 4). We assayed a panel of 10 microsatellite markers for population genetic analyses. Additionally, we constructed and validated a distribution model to generate resistance surfaces for examining the relationship between landscape connectivity, m, θ, and genetic differentiation (FST) using maximum-likelihood population-effects models (MLPE). Populations in continuous habitat were undifferentiated, whereas fragmented populations exhibited genetic structure driven by a single population. Results of the MLPE models in the fragmented landscape revealed spatial variation in θ as the best predictor of pairwise FST, followed by estimates of m, suggesting migration-drift interactions have a stronger influence on genetic differentiation than matrix permeability. Moreover, model coefficients for landscape resistance were comparable between landscapes. Overall, our results provide insight as to how the interaction of gene flow and genetic drift shapes population structure for a dispersal-limited species within a predominately anthropogenic landscape.

Predator avoidance behaviour has been observed in many species and can strongly affect trophic interactions. Animals frequently adjust their response to predators in proportion to risk; however, when they are exposed to chronic predation risk, some animals reduce their antipredator responses in favour of resource acquisition. Animals that are subjected to predation pressure from multiple predator species may have to respond to different spatial and temporal distributions of risk; how they balance these risks will affect their choice of habitat or microhabitat patches. One species that is exposed to a wide range of predators is the pygmy salamander, Desmognathus wrighti. These miniaturized terrestrial plethodontids spend their entire life on the forest floor where they are exposed to a diversity of predators. We examined pygmy salamanders' ability to avoid potential predators based on the presence of substrate-borne chemical cues. The predator species included ground beetles (Carabidae: Pterostichus), Jordan's salamanders, Plethodon jordani, and ring-necked snakes, Diadophis punctatus. The ring-necked snake is a trophic specialist feeding on salamanders while the beetles and Jordan's salamanders are generalist predators. The pygmy salamanders were also exposed to chemical alarm cues from a congeneric salamander (Desmognathus fuscus) and odours from two unfamiliar species: corn snakes (Pantherophis gutattus) and herbivorous desert tortoises (Gopherus agassizii). We used a hierarchical Bayesian beta regression with a repeated measures design to model proportion of time spent in each treatment. Compared to controls, pygmy salamanders exhibited strong aversion to fed ring-necked snakes, a moderate avoidance of unfamiliar odours and a slight attraction to scents from the two common generalist predators. Our results suggest that pygmy salamanders accept modest risk from common generalist predators but avoid less common, specialist predators and unfamiliar species.

Reversible acclimation increases resilience to environmental stress, but acclimation may have hidden costs due to underlying linkages between related physiological traits. These linkages might result in trade‐offs that undermine whole‐organism performance if the change in a related trait reduces the net benefits of acclimation or increases susceptibility to alternative environmental stressors. Metabolic rate and water loss rate are two fundamental physiological traits that could interact due to their dependence on gas exchange across shared physical surfaces. Reductions in water loss rate in response to dehydration stress might reduce metabolic rate by constraining the flux of both water and oxygen. We examined acclimation of metabolic rate and water loss rate using a species of woodland salamander (Plethodon metcalfi) in response to temperature and humidity using a full factorial experimental design. We controlled the evaporative demand of the air across temperatures to assess temperature and humidity as independent cues for acclimation. We predicted that reductions in water loss rate would coincide with reductions in metabolic rate in response to temperature due to shared physical and chemical pathways. We also assessed acclimation of heart rates as a potential compensatory mechanism used to promote oxygen delivery. We integrated these responses into a biophysical model developed from first principles to demonstrate the potential for these interactions to influence habitat suitability. We found that reductions in water loss rates during thermal acclimation were associated with simultaneous reductions in metabolic rates, and we did not find a compensatory response in heart rates. We suggest that these linkages underlie whole‐organism strategies (e.g. physiological dormancy or arousal) for reducing the energetic costs imposed by warm temperatures. The biophysical model suggested that the interaction between these two traits potentially structures phenotypic variation in our population because certain combinations of trait values were incapable of reaching positive energy balance. Trade‐offs between linked physiological traits potentially structure whole‐organism strategies for responding to environmental stressors and constrain phenotypic variation. Therefore, predictions of the benefits of acclimation must be interpreted cautiously without knowledge of the underlying trade‐offs among linked physiological traits. A plain language summary is available for this article.

North America is home to both native and invasive earthworms acting as ecosystem engineers as they build burrows that can serve as habitat for other species or otherwise alter soil structure, affecting nutrient cycling and other ecosystem processes. Here I determine where and what earthworm species commonly occur in my study area, and compare effects of native and invasive earthworms on the common woodland salamander, Plethodon cinereus, in field surveys and laboratory experiments. The native earthworm Eisenoides carolinensis was the most common earthworm, followed by two invasive species Dendrobaena octaedra and Octolasion tyrtaeum. The presence of O. tyrtaeum was associated with a narrower O-horizon (i.e., organic layer in the soil). Using structural equation modeling to explore direct and indirect pathways of these three most common earthworm species on salamanders, I found O. tyrtaeum occurrence was negatively correlated with nighttime salamander counts, a proxy for total salamander numbers, mediated by negative effects on O-horizon depth and microinvertebrate numbers. In the laboratory, O. tyrtaeum and D. octaedra consumed more leaf litter per gram of earthworm per day than the native E. carolinensis. However, salamanders consumed earthworms and used burrows of all native and invasive species of earthworms similarly. The potential for negative indirect effects of the invasive earthworm O. tyrtaeum on P. cinereus was demonstrated both in the field and laboratory, highlighting that seemingly small differences between native and invasive ecosystem engineers have the potential to significantly alter the effects of these closely related native and invasive organisms.

Phenotypic plasticity can be an important adaptive response to climate change, particularly for dispersal-limited species. Temperature frequently alters developmental and phenotypic traits including morphology, behavior, and reproductive cycles. We often lack crucial information about if and how thermal conditions during development will interact with genetic responses and facilitate persistence or adaptation under climate change. Polymorphic species offer an ideal test for this, as alternative morphs often confer differential adaptive advantages. However, few studies have examined the effects of incubation temperature on color expression or development in polymorphic taxa. Here we test if developmental temperature mediates morph frequency in the polymorphic salamander Plethodon cinereus. Although previous research suggests geographic variation in morph proportions results from differential climate adaptation, it remains unknown if plasticity also contributes to this variation. We used a split-clutch common garden experiment to determine the effects of developmental temperature on the color and development of P. cinereus. Our results indicate developmental temperature affects coloration in P. cinereus, either via plasticity or differential mortality, with eggs incubated at warmer temperatures yielding a higher proportion of unstriped individuals than those from cooler temperatures. This temperature response may contribute to the spatial variation in morph frequencies in natural populations. Surprisingly, we found neither temperature nor egg size affected hatchling size. Our study provides important insights into the potential for climate-induced responses to preserve diversity in dispersal-limited species, like P. cinereus, and enable time for adaptive evolution.

Multiple pathways exist for species to respond to changing climates. However, responses of dispersal‐limited species will be more strongly tied to ability to adapt within existing populations as rates of environmental change will likely exceed movement rates. Here, we assess adaptive capacity in Plethodon cinereus, a dispersal‐limited woodland salamander. We quantify plasticity in behavior and variation in demography to observed variation in environmental variables over a 5‐year period. We found strong evidence that temperature and rainfall influence P. cinereus surface presence, indicating changes in climate are likely to affect seasonal activity patterns. We also found that warmer summer temperatures reduced individual growth rates into the autumn, which is likely to have negative demographic consequences. Reduced growth rates may delay reproductive maturity and lead to reductions in size‐specific fecundity, potentially reducing population‐level persistence. To better understand within‐population variability in responses, we examined differences between two common color morphs. Previous evidence suggests that the color polymorphism may be linked to physiological differences in heat and moisture tolerance. We found only moderate support for morph‐specific differences for the relationship between individual growth and temperature. Measuring environmental sensitivity to climatic variability is the first step in predicting species' responses to climate change. Our results suggest phenological shifts and changes in growth rates are likely responses under scenarios where further warming occurs, and we discuss possible adaptive strategies for resulting selective pressures.

Prescribed fire is used in the central Appalachians to promote and maintain mixed-oak and pine forests, create open forest conditions, improve habitat for wildlife, and to reduce the risk of impact of higher intensity wildfires on human development. Few studies have investigated responses of terrestrial salamander populations to habitat management using fire, and estimated responses have been neutral, negative, and positive depending on geography, species, and fire-severity. We examined woodland salamander (genus Plethodon) population responses to habitat management using prescribed fire on Shenandoah Mountain in the George Washington National Forest in West Virginia and Virginia, USA. We focused on responses of the Cow Knob Salamander (P. punctatus), a talus specialist and species of high conservation concern, but also examined responses of the Eastern Red-Backed Salamander (P. cinereus), a widespread habitat generalist. Three burn units were subjected to two low-severity burns and one unit was burned five times with ca. 40% tree mortality. Using a combination of nighttime visual encounter surveys and coverboard surveys, we compared terrestrial salamander abundance and body condition in unburned and burned areas. We also measured habitat characteristics at sampling sites to determine if prescribed burn histories were correlated with habitat conditions important to woodland salamanders. Mean abundance for P. punctatus was lower at sites that were burned, but there was not a strong burn effect for P. cinereus. Abundance of both species was positively correlated with canopy cover. Mean and median body condition index (BCI) score was higher for P. punctatus and lower for P. cinereus on the West Virginia side of Shenandoah Mountain, and lower in burned areas for both species. However, the most parsimonious BCI models did not contain the burn predictor. Management using prescribed fire altered microhabitat conditions that are important for woodland salamanders, such as canopy cover, leaf litter depth, and vegetation groundcover. Our study suggests that woodland salamanders in the central Appalachians can persist in forests managed using prescribed fire, but also indicates that prescribed fire can result in reduced habitat quality for some woodland salamander species.

We examined responses of the woodland salamander Ensatina ( Ensatina eschscholtzii ) to commercial thinning by helicopter in late-seral Douglas-fir forest in northwestern California, USA, using a before-after control-impact (BACI) design. We employed passive pitfall traps on eight (four each treatment and control) 100 trap grids over eight years (1996–2003), three years pre- and five years post-treatment. We conducted 3–11 weekly site visits within each of 10 trap periods depending upon weather over two spring and eight fall periods for a total of 50 visits to test for differences in surface counts and body condition. We standardized data to captures/1000 trap nights. Surface counts over the eight years were highly variable with less surface activity in spring compared to fall. Although the BACI design allows for initial differences among pre-treatment grids there were none. However, in the three years post-treatment (1999–2001) mean counts on treatment grids were significantly lower than on control grids. By 2002 and 2003, post-treatment counts on control and treatment grids both declined substantially, such that despite 47% more salamanders on control grids, variability was sufficient to preclude statistical significance. Recapture rates averaged 23.3% for three years pre-treatment, but increased to 51.4% for the five years post-treatment. Pre-treatment population estimates increased on control grids post-treatment, however, surface density estimates declined. On treatment grids both population and density estimates declined following treatment. Body condition analyses indicated no significant differences between controls and treatments for either sex prior to or post-treatment. The three years of significant differences in surface counts between controls and treatments post-treatment, followed by no differences in the last two years of the study, combined with no differences in BCIs, lead us to conclude that the surface activity of Ensatina on treatments grids was only temporarily impacted by the thinning. Declining population estimates on treatment grids also indicated at least a temporary reduction in those populations. We unexpectedly found declines in counts and density estimates on both sets of grids over the eight years, suggesting a wider effect may be causing reductions in the surface activity of these salamanders, and possibly in their populations.

Diverse bacteria inhabit amphibian skin; some of those bacteria inhibit growth of the fungal pathogen Batrachochytrium dendrobatidis . Yet there has been no systematic survey of anti- B. dendrobatidis bacteria across localities, species, and elevations. This is important given geographic and taxonomic variations in amphibian susceptibility to B. dendrobatidis . Our collection sites were at locations within the Appalachian Mountains where previous sampling had indicated low B. dendrobatidis prevalence. We determined the numbers and identities of anti- B. dendrobatidis bacteria on 61 Plethodon salamanders (37 P. cinereus , 15 P. glutinosus , 9 P. cylindraceus ) via culturing methods and 16S rRNA gene sequencing. We sampled co-occurring species at three localities and sampled P. cinereus along an elevational gradient (700 to 1,000 meters above sea level [masl]) at one locality. We identified 50 anti- B. dendrobatidis bacterial operational taxonomic units (OTUs) and found that the degree of B. dendrobatidis inhibition was not correlated with relatedness. Five anti- B. dendrobatidis bacterial strains occurred on multiple amphibian species at multiple localities, but none were shared among all species and localities. The prevalence of anti- B. dendrobatidis bacteria was higher at Shenandoah National Park (NP), VA, with 96% (25/26) of salamanders hosting at least one anti- B. dendrobatidis bacterial species compared to 50% (7/14) at Catoctin Mountain Park (MP), MD, and 38% (8/21) at Mt. Rogers National Recreation Area (NRA), VA. At the individual level, salamanders at Shenandoah NP had more anti- B. dendrobatidis bacteria per individual (μ = 3.3) than those at Catoctin MP (μ = 0.8) and at Mt. Rogers NRA (μ = 0.4). All salamanders tested negative for B. dendrobatidis . Anti- B. dendrobatidis bacterial species are diverse in central Appalachian Plethodon salamanders, and their distribution varied geographically. The antifungal bacterial species that we identified may play a protective role for these salamanders. IMPORTANCE Amphibians harbor skin bacteria that can kill an amphibian fungal pathogen, Batrachochytrium dendrobatidis . Some amphibians die from B. dendrobatidis infection, whereas others do not. The bacteria that can kill B. dendrobatidis , called anti- B. dendrobatidis bacteria, are thought to influence the B. dendrobatidis infection outcome for the amphibian. Yet how anti- B. dendrobatidis bacterial species vary among amphibian species and populations is unknown. We determined the distribution of anti- B. dendrobatidis bacterial species among three salamander species ( n = 61) sampled at three localities. We identified 50 unique anti- B. dendrobatidis bacterial species and found that all of the tested salamanders were negative for B. dendrobatidis . Five anti- B. dendrobatidis bacterial species were commonly detected, suggesting a stable, functional association with these salamanders. The number of anti- B. dendrobatidis bacteria per individual varied among localities but not among co-occurring salamander species, demonstrating that environment is more influential than host factors in structuring the anti- B. dendrobatidis bacterial community. These anti- B. dendrobatidis bacteria may serve a protective function for their salamander hosts.

Evolutionary change has been demonstrated to occur rapidly in human‐modified systems, yet understanding how multiple components of global change interact to affect adaptive evolution remains a critical knowledge gap. Climate change is predicted to impose directional selection on traits to reduce thermal stress, but the strength of directional selection may be mediated by changes in the thermal environment driven by land use. We examined how regional climatic conditions and land use interact to affect genetically based color polymorphism in the eastern red‐backed salamander (Plethodon cinereus). P. cinereus is a woodland salamander with two primary discrete color morphs (striped, unstriped) that have been associated with macroclimatic conditions. Striped individuals are most common in colder regions, but morph frequencies can be variable within climate zones. We used path analysis to analyze morph frequencies among 238,591 individual salamanders across 1,170 sites in North America. Frequency of striped individuals was positively related to forest cover in populations occurring in warmer regions (>7°C annually), a relationship that was weak to nonexistent in populations located in colder regions (≤7°C annually). Our results suggest that directional selection imposed by climate warming at a regional scale may be amplified by forest loss and suppressed by forest persistence, with a mediating effect of land use that varies geographically. Our work highlights how the complex interaction of selection pressures imposed by different components of global change may lead to divergent evolutionary trajectories among populations.

The Woodland Salamander genus Plethodon Tschudi, 1838, consists of terrestrial lungless salamanders, has been cited as an exemplar of a lineage characterized by niche conservatism. This conclusion has contributed to broader hypotheses dealing with the role of niche conservatism in speciation and the maintenance of species diversity. We re-evaluated these salamanders using newly published techniques designed to detect niche conservatism versus niche divergence using computer-based niche modeling and spatial analysis within a phylogenetic framework. We specifically studied parapatric neighbors in the species complex of Plethodon glutinosus (Green, 1818) to determine if there is evidence of a role for niche divergence at speciation or if niche conservatism characterizes the complex as has been reported for the larger genus. We found that new statistical approaches yield different results from earlier work, suggesting that niche divergence has been a much more important player in the speciation process than has heretofore been understood. Although different parts of the overall Plethodon phylogeny may be characterized by different rates of niche evolution, the rapidly burgeoning area of research centered on niche conservatism appears to be in a state of flux with regards to methods that give consistently repeatable results.

In addition to creating or modifying habitat, ecosystem engineers interact with other species as predators, prey, or competitors. The earthworm, Lumbricus terrestris, interacts with the common woodland salamander, Plethodon cinereus, via: (1) ecosystem engineering, by providing burrows that are used as a refuge, (2) direct effects as a prey item, and (3) indirectly, by competing with microinvertebrates, another prey item for P. cinereus. Using enclosures in the forest, I examined the relative strengths of these component pathways between seasons and salamander age classes. I found that the relative strength (partial eta2) of the positive direct (trophic) effect of L. terrestris on the change in mass of P. cineresus was greater than that of the negative indirect effect, but only in summer. Positive effects of ecosystem engineering were only evident over the winter as increased adult survival. This research has implications for how habitat provisioning complements more well-studied species interactions, such as competition and predation, within communities.

In the competition to acquire mates, environmental factors can be important in determining the relative quality of an individual. These aspects of quality are often conveyed through signals used for mate assessment by the most energetically invested sex. In red-backed salamanders, Plethodon cinereus, chemical signals communicate a surprising amount of information, much of which is dependent on the current condition of the sender. One such condition-dependent aspect of information conveyed via chemicals is the quality of the sender’s diet, but it is unclear as to whether this information is actively advertised by the sender (i.e., a signal) or simply inferred from food-derived odors (i.e., a cue). The amount of time on different diets required for changes in signaling is also unknown. I examined how quickly gravid female salamanders could detect a difference between the scents of males on high- vs. low-quality diets without fecal cues. The amount of protein present in two known signaling glands (the mental and postcloacal glands) was also measured after experimental feeding. Gravid females were able to infer the differences in male diet quality after only 1 week. Females also responded to the male scents more quickly after differential feeding had begun. High-quality males had significantly more protein present in both signaling glands than low-quality males. This scenario highlights the ongoing interplay between the quality of an individual and its environment, with males actually advertising the status of this relationship as an honest signal for mate assessment.

Using coverboard arrays, we monitored woodland salamanders on the Fernow Experimental Forest in the central Appalachian Mountains, West Virginia, USA prior to and following two prescribed fires in mixed oak (Quercus spp.) forest stands. Treatments were burn plots on upper slopes or lower slopes fenced to prevent white-tailed deer (Odocoileus virginianus) herbivory or control plots that were unfenced and unburned. Most of the 7 species we observed were the mountain dusky salamander (Desmognathus ocropheaus), red-backed salamander (Plethodon cinereus) and slimy salamander (Plethodon glutinosis). Significant population responses were difficult to interpret with numerous treatment and year interactions. Results largely were equivocal. We found no change in woodland salamander assemblage prior to burning or afterwards. There were few differences in adult to juvenile ratios of salamanders among treatments. Still, a priori contrasts of mountain dusky salamanders and red-backed salamander counts corrected for detection probability were greater under coverboards in the 2 years monitored after both prescribed fires had occurred than before burning or in unburned controls. This suggests that these species responded to the reduced leaf litter on the forest floor by utilizing coverboards more. Similarly, the three predominate species of salamanders also were more numerous under coverboards in plots subjected to deer herbivory with less subsequent forest floor vegetation as compared to those burned plots that were fenced. Our observations would suggest that woodland salamanders somewhat are tolerant of two prescribed fires within close temporal proximity. However, because woodland salamanders can be significantly reduced following timber harvest, continued research is needed to fully understand impacts of fire as a pre-harvest management tool in central Appalachian forests.

Studies examining the influence of habitat provisioning by one species on the behavior of other species can provide key insights regarding impacts of ecosystem engineers on the availability of resources to other species. More specifically, an organism’s use of additional habitat provided by ecosystem engineers may affect the interpretation of observational or demographic data. We chose to examine the possible influence of earthworms, as ecosystem engineers, on the behavior of terrestrial salamanders, common forest vertebrates in North America. We conducted two experiments in microcosms to examine whether the earthworm Lumbricus terrestris acted as an ecosystem engineer by providing additional habitat (burrows) for Plethodon cinereus, a common woodland salamander. We also examined whether the behavioral changes of the non-burrowing, fossorial P. cinereus differed across age classes. Adults and juveniles responded similarly, with both adults and juveniles found under cover objects more often when earthworms were absent and using earthworm burrows when available. Our field data supported our experimental data in that greater numbers of earthworms on field plots were associated with a lower proportion of cover objects being occupied by salamanders (i.e., salamanders were likely underground). We found no differences in the response of adults and juveniles to the presence of earthworms. By providing underground habitat for a common terrestrial amphibian, the presence of earthworms may impact demographic studies on salamanders and conclusions drawn from those studies regarding aboveground faunal communities in eastern North America.

Woodland salamander responses to either traditional grazing or silvopasture systems are virtually unknown. An information-theoretic modelling approach was used to evaluate responses of red-backed salamanders (Plethodon cinereus) to silvopasture and meadow conversions in southern West Virginia. Searches of area-constrained plots and artificial coverboards that were distributed across a gradient of agricultural conversion and grazing intensity, including hardwood silvopastures, hay meadows, forest edges, and reference forests yielded 2823 salamanders between May 2004 and November 2005. Salamander presence and abundance were positively associated with increasing cover of herbaceous vegetation and negatively associated with the intensity of agricultural disturbance. Although salamander presence and abundance appeared to be negatively influenced by agricultural disturbance when compared to reference forest conditions, the occurrence of red-backed salamanders within agriculturally modified habitats indicates this species may be more resilient to forest conversion than previously thought. This study suggests that herbaceous vegetation retained within some agricultural treatments, in combination with artificial cover, may at least partially mitigate the loss of forest canopy for red-backed salamanders.

The ability of animal species to disperse through matrix habitats is likely to have important implications for species' responses to habitat fragmentation. Within fragmented forests, open fields are among the most common matrix habitats. However, few empirical studies have measured the effects of open habitats on dispersal success of forest-dwelling species. We used two experiments to determine the effects of open habitats and distance on dispersal and colonization by red-backed salamanders, Plethodon cinereus. In the first experiment, salamanders were displaced from habitat edges 25 m and 55 m into an open field and farther into the forest. Return rates from salamanders crossing the field vs. the forest were then used to determine the barrier effects of open habitats. Although return rate declined with distance, salamanders returned successfully through open habitat as often as they returned successfully through forest. In the second experiment, we constructed “islands” of forest-like habitat within an o...

Aggressive interference can be an important expression of competitive interactions within and among species. We studied potential interference between two similar-sized, sympatric species of woodland salamander. The Pigeon Mountain Salamander, Plethodon petraeus, occurs in habitat patches of rocky outcroppings along a restricted area on the eastern flank of Pigeon Mountain in northwestern Georgia. This species is relatively rare in the intervening areas of forest-floor habitat, which is occupied by a similar-sized, potential competitor, the Slimy Salamander (Plethodon glutinosus). Both species exhibited aggression and defended territories against conspecifics and heterospecifics in laboratory-based encounters. Individuals of P. glutinosus were superior in aggressive encounters with individuals of P. petraeus. Coexistence of the two species occurs because habitats are partially exclusive. There is enough habitat overlap, however, to generate possible interspecific competition. We hypothesize that the presence of territorial, aggressive P. glutinosus in areas between habitat patches may influence interpatch dispersal by P. petraeus.

Recent concern over global amphibian population declines has highlighted a need for more extensive, rigorous monitoring programs. Two sources of variation, spatial variation and variation in detection probability, make the design and implementation of effective monitoring programs difficult. We used Pollock's robust design in a 3-year capture–recapture study to estimate detection probability and temporary emigration for Plethodon salamanders in Great Smoky Mountains National Park (Tennessee/North Carolina), USA. We used 12 competing models to determine the importance of temporary emigration, and we explored temporal and behavioral effects on conditional capture probabilities. The top 4 models all included random temporary emigration, and Akaike model weights indicated that this parameter was the most important. Models that contained behavioral effects in capture probabilities were selected more often than models with equal capture probabilities for marked and previously unmarked individuals. The ...

Guilds consist of groups of species that use mutual resources or collections of overlapping resources. Guilds are often comprised of members of the same family or genus, but they can span larger taxonomic categories. Predacious arthropods, such as carabid beetles, centipedes and spiders, and small carnivorous vertebrates, such as woodland salamanders, overlap extensively in microhabitat and prey types. These taxa, though unrelated, may form important guilds of forest floor predators. We examined the behavioral interactions between a carabid ground beetle (Platynus tenuicollis) and a small woodland salamander (the red-backed salamander, Plethodon cinereus) in laboratory encounters. These species occur syntopically and feed on similar prey. Individuals of P. cinereus are territorial and defend cover objects against conspecific and congeneric intruders. Because both species require moist conditions, it seems probable that they would compete for resources, such as cover objects and prey, as the fores...

We surveyed cove hardwood stands aged 15, 25, 50, and ≥85 years following clearcutting in the southern Appalachian Mountains of northern Georgia to assess the effects of stand age and stand habitat characteristics on salamander communities using drift-fence array and pitfall methodologies from May 1994 to April 1995. Over a 60,060 pitfall trapnight effort, we collected 3937 salamanders represented by Desmognathus aeneus, Desmognathus monticola, Desmognathus ocoee, Desmognathus quadramaculatus, Eurycea bislineata, Gyrinophilus porphyriticus, Pseudotriton ruber, Plethodon glutinosus, Plethodon serratus, and Notophthalmus viridescens. Analysis of covariance with pitfall array to stream distance as the covariate showed that salamander species richness and diversity measures and numbers of Desmognathus aeneus and Desmognathus ocoee were highest in stands ≥85 years. Eurycea bislineata and Plethodon glutinosus were more abundant in stands ≤50 years old than in stands ≥85 years. Within cove hardwood stands, species richness and diversity measures and relative abundances of Desmognathus spp. and Gyrinophilus porphyriticus were negatively correlated with distance to stream. Species richness and diversity were positively correlated to amounts of emergent rock. Species richness, diversity and relative abundances of Desmognathus spp. were correlated with basal area within stands and extent of connected mesic, cove hardwood habitat and amount of cove habitat within 1 km radius among stands. Eurycea bislineata was negatively correlated with landform index, a measure of surrounding landform sheltering, and Plethodon glutinosus was positively correlated with elevation in cove hardwood stands. Our research indicates stand age is an important factor in explaining the abundance and community composition of salamanders in southern Appalachian cove hardwood communities. Because southern Appalachian woodland salamander communities are slow to recover and are substantially changed following disturbances such as clearcutting, populations in small, isolated cove hardwood stands might be more vulnerable to extirpation or may require longer recovery times than those in larger coves. Managers may need to assess habitat features such as cove extent and habitat connectivity to minimize impacts on these taxa by forest management activities in southern Appalachian cove hardwood communities.

Landscape genetics has seen tremendous advances since its introduction, but parameterization and optimization of resistance surfaces still poses significant challenges. Despite increased availability and resolution of spatial data, few studies have integrated empirical data to directly represent ecological processes as genetic resistance surfaces. In our study, we determine the landscape and ecological factors affecting gene flow in the western slimy salamander (Plethodon albagula). We used field data to derive resistance surfaces representing salamander abundance and rate of water loss through combinations of canopy cover, topographic wetness, topographic position, solar exposure and distance from ravine. These ecologically explicit composite surfaces directly represent an ecological process or physiological limitation of our organism. Using generalized linear mixed-effects models, we optimized resistance surfaces using a nonlinear optimization algorithm to minimize model AIC. We found clear support for the resistance surface representing the rate of water loss experienced by adult salamanders in the summer. Resistance was lowest at intermediate levels of water loss and higher when the rate of water loss was predicted to be low or high. This pattern may arise from the compensatory movement behaviour of salamanders through suboptimal habitat, but also reflects the physiological limitations of salamanders and their sensitivity to extreme environmental conditions. Our study demonstrates that composite representations of ecologically explicit processes can provide novel insight and can better explain genetic differentiation than ecologically implicit landscape resistance surfaces. Additionally, our study underscores the fact that spatial estimates of habitat suitability or abundance may not serve as adequate proxies for describing gene flow, as predicted abundance was a poor predictor of genetic differentiation.

Herpetofauna responses to forest management and early successional habitats are influenced by species-specific adaptations to historical disturbance regimes. It can take decades for woodland salamander diversity to recover after heavy overstory removal for even-aged forest regeneration or hot fires that yield higher light, drier microclimates, and reduced leaf litter cover, but some frog and toad species may tolerate or even increase after disturbances. In particular, disturbances that retain some canopy cover, such as selection harvests or low intensity burns, can mitigate effects on terrestrial salamanders. The same early successional conditions that are detrimental to salamanders can benefit many reptile species, such as fence lizards (Sceloporus undulatus). Maintaining stand age diversity across central hardwood forest landscapes, including retention of mature forest communities, should provide habitats for both early successional wildlife and mature forest species.

We compared the species richness and abundance of salamanders on six recent clearcuts ( 50 years old) in southern Appalachian forests in western North Carolina, U.S.A. Catches of salamanders from plots in mature forest stands were about five times higher than those on recent clearcuts. Almost all species and major taxonomic groups of salamanders were adversely affected by timber removal. Mean number of species collected per plot was about twice as great in mature forest stands as in clearcuts. Analyses of stand age versus salamander catch for 47 plots indicate that 50–70 years are required for populations to return to predisturbance levels following cutting. We conservatively estimate that clearcutting in U.S. national forests in western North Carolina results in a loss of nearly 14 millton salamanders annually. It also is chronically reducing regional populations by more than a quarter of a billion salamanders (9%) below that which could be sustained if mature forests were not cut. Nosotros comparamos la riqueza de especies y abundancia de las salamandras en seis recientes cortas totales de bosque ( 50 anos) en los bosques del sur de los Apalaches, en el oeste de Carolina del Norte, Estados Unidos. La captura de salamandras por plot en bosques de rodales maduros fue aproximadamente cinco veces mayor que en aquellos cortados recientemente. Casi todas las especies y los mayores grupos taxonomicos de salamandras fueron adversamente afectados por la tala. La media del numero de especies capturada por plot fue aproximadamente dos veces mayor en los rodales de bosques maduros que en los cortados recientemente. Los analisis de edad del rodal versus captura de salamandras para 47 plots indican que entre 50–70 anos son necesarios para que las poblaciones retornen a los niveles previos a la perturbacion ocasionada por la tala del bosque. Nosotros estimamos en una forma conservadora que la tala de los Bosques Nacionales en el oeste de Carolina del Norte trae como consecuencia una perdida de alrededor de 14 millones de salamandras anualmente. La tala tambien esta reduciendo cronicamente las poblaciones a nivel regional en mas de un cuarto de billones de salamandras (9%) por debajo del nivel que podria ser sostenido por los bosques maduros si no hubieran sido cortados.

Ecosystem engineers create habitat that can be used by other species in multiple ways, such as refuges from predators, places to breed, or areas with increased prey resources. I conducted a series of enclosure experiments to: (1) determine if salamanders use earthworm burrows, and (2) examine the potential influence of earthworm burrow use and indirect effects on salamander intra- and interspecific competition, predator avoidance, and seasonal performance. I found that one species of woodland salamander, Plethodon cinereus, used earthworm burrows 50% of the time when burrows were present. Neither adults nor juveniles of the congeneric P. glutinosus used earthworm burrows. Intraspecific, but not interspecific, competition by P. cinereus affected salamander behavior when earthworms were absent, with P. cinereus found under cover objects >70% of the time when alone or with a P. glutinosus, but only 40% of the time when with another P. cinereus. When earthworms were present, the behavior of P. cinereus was similar across salamander treatments. Earthworms decreased the amount of leaf litter and microinvertebrates, although this did not affect salamander mass. In subsequent experiments using only P. cinereus, the refuge provided by earthworm burrows increased the survival of P. cinereus over the winter and allowed P. cinereus to avoid being consumed by the common garter snake (Thamnophis sirtalis). Because earthworm burrows provide a refuge for P. cinereus during intraspecific encounters, in the presence of a predator and over the winter, they may serve as an important belowground–aboveground linkage in eastern forests where salamanders are common.

Factors that negatively affect the quality of wildlife habitat are a major concern for conservation. Non-native species invasions, in particular, are perceived as a global threat to the quality of wildlife habitat. Recent evidence indicates that some changes to understory plant communities in northern temperate forests of North America, including invasions by 3 non-native plant species, are facilitated by non-native earthworm invasion. Furthermore, non-native earthworm invasions cause a reduction in leaf litter on the forest floor, and the loss of forest leaf litter is commonly associated with declines in forest fauna, including amphibians. We conducted a mark-recapture study of woodland salamander abundance across plant invasion fronts at 10 sites to determine whether earthworm or plant invasions were associated with reduced salamander abundance. Salamander abundance declined exponentially with decreasing leaf litter volume. There was no significant relationship between invasive plant cover and salamander abundance, independent of the effects of leaf litter loss due to earthworm invasion. An analysis of selected salamander prey abundance (excluding earthworms) at 4 sites showed that prey abundance declined with declining leaf litter. The loss of leaf litter layers due to non-native earthworm invasions appears to be negatively affecting woodland salamander abundance, in part, because of declines in the abundance of small arthropods that are a stable resource for salamanders. Our results demonstrate that earthworm invasions pose a significant threat to woodland amphibian fauna in the northeastern United States, and that plant invasions are symptomatic of degraded amphibian habitat but are not necessarily drivers of habitat degradation.

We examined local and landscape-scale variable influence on the depth and magnitude of edge effects on woodland salamanders in mature mixed mesophytic and northern hardwood forest adjacent to natural gas well sites maintained as wildlife openings. We surveyed woodland salamander occurrence from June–August 2006 at 33 gas well sites in the Monongahela National Forest, West Virginia. We used an information-theoretic approach to test nine a priori models explaining landscape-scale effects on woodland salamander capture proportion within 20 m of field edge. Salamander capture proportion was greater within 0–60 m than 61–100 m of field edges. Similarly, available coarse woody debris proportion was greater within 0–60 m than 61–100 m of field edge. Our ASPECT model, that incorporated the single variable aspect, received the strongest support for explaining landscape-scale effects on salamander capture proportion within 20 m of opening edge. The ASPECT model indicated that fewer salamanders occurred within 20 m of opening edges on drier, hotter southwestern aspects than in moister, cooler northeastern aspects. Our results suggest that forest habitat adjacent to maintained edges and with sufficient cover still can provide suitable habitat for woodland salamander species in central Appalachian mixed mesophytic and northern hardwood forests. Additionally, our modeling results support the contention that edge effects are more severe on southwesterly aspects. These results underscore the importance of distinguishing among different edge types as well as placing survey locations within a landscape context when investigating edge impacts on woodland salamanders.

Several studies have reported climate-associated changes in phenotypically plastic traits of amphibians, yet it remains unknown whether amphibians can manifest an evolutionary response to global climate change at the rate and magnitude that it is occurring. To assess this issue, we examined temporal change in the morphology of the red-backed salamander (Plethodon cinereus), a small, abundant woodland salamander distributed widely in eastern North America with two distinct morphotypes: striped individuals associated with cooler microclimates and unstriped individuals associated with warmer microclimates. We compiled morph frequencies for 50,960 individual salamanders from 558 sites as recorded in the published literature and in unpublished field notes of herpetologists between 1908 and 2004. We observed that striping probability increased with increasing latitude, longitude, and elevation and decreased (from 80% to 74% range wide) with time. The combined forces of regional climate warming and, particularly, forest disturbance have evidently been sufficient to cause morphological evolution in this amphibian over the last century.

The natural density of red-backed salamanders, Plethodon cinereus, is 4–5 times greater at the New York study site in hardwood than in hemlock forests. The hypothesis that physico-chemical differences between forest types differentially affect the physiology of P. cinereus was tested. Salamanders were confined in individual enclosures in either a beech or a hemlock forest for 22 d. At the end of the enclosure period, they were analyzed for total body content of sodium, potassium, calcium, and magnesium, and for wet mass, dry mass, and body water concentration. Salamanders from the two forest types did not differ in any body parameter after the 22 d enclosure period. Also, the hypothesis that artificially acidified “rain” would inhibit the ability of salamanders to maintain ion balance was tested. Salamanders were confined for 34 d in individual enclosures in the two forests and exposed periodically to artificial rain of either pH 3 or pH 5. Soil pHs in the enclosures changed during the enclosure period, and artificial rain of both pHs caused soil pHs in the hemlock forest to increase even though pHs outside enclosures decreased. Enclosures that received artificial rain of pH 3 had lower soil pHs than enclosures that received artificial rain of pH 5. Salamanders in enclosures at pH 3 had reduced body sodium levels and tended to have lower body water concentration than salamanders in enclosures at pH 5 or controls. Thus, these terrestrial salamanders in a field setting demonstrated a physiological response to simulated acidic deposition consistent with the effects of exposure to acidic substrate previously observed in the laboratory. Since P. cinereus is frequently an abundant member of forest floor communities, and since it is sensitive to altered chemical inputs, it may be a useful indicator for assessing anthropogenic changes in ecosystems.

Using museum collection records and variables computed by digital terrain modeling in a geographic information system, we examined the relationship of elevation, aspect, and “cove” patch size to the presence or absence of 7 common woodland salamanders in mature cove hardwood and northern hardwood forests in the southern Appalachians of Georgia, North Carolina, and South Carolina. Overall, elevation, aspect and patch size were poor discriminators among salamander species presence or absence at collection sites. Increased elevation was an important variable explaining the presence of Ocoee salamanders (Desmognathus ocoee) and Jordan's salamanders (Plethodon jordani). In contrast, decreased elevation was an important variable explaining the presence of slimy salamanders (Plethodon glutinosus). Our study contrasts with recent research indicating that suitable habitat patch size is an important determinant of woodland salamander species richness and abundance at recently disturbed sites. In these matu...