Plethodon Cinereus Research Articles

Reducing uncertainty with iterative model updating parses effects of competition and environment on salamander occupancy.

Making timely management decisions is often hindered by uncertainty. Monitoring reduces two key types of uncertainty. First, it serves to reduce structural uncertainty of how the system works and provides support for expectations of how a system works. Second, it serves to reduce parametric uncertainty of the drivers of system dynamics. By combining monitoring data and quantitative models, we can reduce structural and parametric uncertainty. To demonstrate this, we focus on the Shenandoah salamander (Plethodon shenandoah), a United States Federally Endangered Species. Early work suggested that P. shenandoah extinction risk results from competition with a conspecific (Plethodon cinereus). However, more recent work has found equivocal support for this claim, instead suggesting that abiotic factors, such as moisture and temperature, drive P. shenandoah persistence. Using long-term monitoring data, we find that while competition may play a part in P. shenandoah extinction risk, measures of surface moisture are better predictors of occupancy dynamics. Further, we find decreased detection rates of P. shenandoah when P. cinereus is present, suggesting a conflation of detection probability with actual competition, which cautions against making inference from unadjusted observations of occurrence. Using multiple lines of inquiry allows for more robust understanding of system drivers in the face of high uncertainty, increasing opportunities to manage extinction risk.

The Limits of the Metapopulation: Lineage Fragmentation in a Widespread Terrestrial Salamander (Plethodon cinereus)

Abstract In vicariant species formation, divergence results primarily from periods of allopatry and restricted gene flow. Widespread species harboring differentiated, geographically distinct sublineages offer a window into what may be a common mode of species formation, whereby a species originates, spreads across the landscape, then fragments into multiple units. However, incipient lineages usually lack reproductive barriers that prevent their fusion upon secondary contact, blurring the boundaries between a single, large metapopulation-level lineage and multiple independent species. Here, we explore this model of species formation in the Eastern Red-backed Salamander (Plethodon cinereus), a widespread terrestrial vertebrate with at least 6 divergent mitochondrial clades throughout its range. Using anchored hybrid enrichment data, we applied phylogenomic and population genomic approaches to investigate patterns of divergence, gene flow, and secondary contact. Genomic data broadly match most mitochondrial groups but reveal mitochondrial introgression and extensive admixture at several contact zones. While species delimitation analyses in Bayesian Phylogenetics and Phylogeography supported 5 lineages of P. cinereus, genealogical divergence indices (gdi) were highly sensitive to the inclusion of admixed samples and the geographic representation of candidate species, with increasing support for multiple species when removing admixed samples or limiting sampling to a single locality per group. An analysis of morphometric data revealed differences in body size and limb proportions among groups, with a reduction of forelimb length among warmer and drier localities consistent with increased fossoriality. We conclude that P. cinereus is a single species, but one with highly structured component lineages of various degrees of independence.

Methylmercury Concentrations More Strongly Associated With Trait Variation Than Food Web Position in Plethodontid Salamanders.

Salamanders serve as bioindicators of mercury (Hg) in both terrestrial and aquatic habitats and are an important link in the food web between low-trophic prey and higher-trophic predators. We investigated the drivers of methylmercury (MeHg) exposure in three common plethodontid salamander species in New York State, USA, including comparisons among regions, habitat types (terrestrial and semiaquatic), and color morphs of Plethodon cinereus (striped and unstriped). Nonlethal tail samples were collected from one terrestrial species (P. cinereus) and two semiaquatic species (Eurycea bislineata and Desmognathus spp.) in the Adirondack Mountains (ADK) and the Finger Lakes National Forest (FLNF) regions. Samples were analyzed for MeHg and stable isotopes, including δ15N and δ13C which are proxies of trophic position and diet, respectively. Despite elevated biota Hg concentrations typically found in the ADK, salamander MeHg concentrations did not differ by region in the terrestrial species and one of the semiaquatic species. In addition, diet and trophic level did not explain MeHg exposure in salamanders. Semiaquatic salamanders exhibited higher MeHg concentrations than terrestrial salamanders in the FLNF only. Within species, only snout-vent length predicted MeHg concentrations in E. bislineata with few other variables significant as predictors of MeHg concentrations in path models. Among P. cinereus individuals in the FLNF, the striped morph had greater MeHg concentrations than the unstriped morph, and food web tracers were not different between morphs. Overall, New York State salamander Hg concentrations were elevated compared to other locations where these species are present. The present study establishes baseline Hg data in salamanders for future assessments of changes in Hg bioavailability to forests of New York State. Environ Toxicol Chem 2024;43:2045-2057. © 2024 SETAC.

Range-wide salamander densities reveal a key component of terrestrial vertebrate biomass in eastern North American forests.

Characterizing the population density of species is a central interest in ecology. Eastern North America is the global hotspot for biodiversity of plethodontid salamanders, an inconspicuous component of terrestrial vertebrate communities, and among the most widespread is the eastern red-backed salamander, Plethodon cinereus. Previous work suggests population densities are high with significant geographic variation, but comparisons among locations are challenged by lack of standardization of methods and failure to accommodate imperfect detection. We present results from a large-scale research network that accounts for detection uncertainty using systematic survey protocols and robust statistical models. We analysed mark-recapture data from 18 study areas across much of the species range. Estimated salamander densities ranged from 1950 to 34 300 salamanders ha-1, with a median of 9965 salamanders ha-1. We compared these results to previous estimates for P. cinereus and other abundant terrestrial vertebrates. We demonstrate that overall the biomass of P. cinereus, a secondary consumer, is of similar or greater magnitude to widespread primary consumers such as white-tailed deer (Odocoileus virginianus) and Peromyscus mice, and two to three orders of magnitude greater than common secondary consumer species. Our results add empirical evidence that P. cinereus, and amphibians in general, are an outsized component of terrestrial vertebrate communities in temperate ecosystems.

Evaluating elements of artificial cover object design for terrestrial salamander monitoring

AbstractDeclines in amphibian populations have led to greater research and monitoring of species of interest, including terrestrial salamanders. Several methods have been used to study salamanders, including artificial cover objects (ACOs). Artificial cover objects are easy to install and use, limit destruction to habitats, are easily reproducible both across and within studies, and can reduce variability between observers and sites by not relying on observer search time or effort, or on natural cover. However, different ACO designs and spacing may yield different capture rates, and thus different estimates of population densities. We examined the influence of ACO wood type, size, shape, and spacing arrays on the number of individually marked red‐backed salamanders (Plethodon cinereus) captured under ACOs. White pine (Pinus strobus) ACOs yielded the most (average of 14 across 3 replicates) individuals, while white ash (Fraxinus americana) ACOs had the least (average of 7 across 3 replicates). Larger ACOs had similar capture rates compared to equal sampled areas of smaller ACOs, but a single smaller ACO tended to have fewer individual salamander captures than a single large ACO. Our results suggested that high‐density ACO spacing arrays may increase capture probability but reduce the area uniquely sampled by each ACO. In contrast, low‐density ACO arrays may produce population estimates more comparable to other arrays. Future studies should consider monitoring goals and available sampling effort when choosing a spacing array. As ACO study design impacts salamander capture rates, a clear understanding of the effects of each element of ACO design is important when drawing inferences or making comparisons among studies.

Updated range map of an endangered salamander and congeneric competitor reveals different niche preferences.

Estimating distributions for cryptic and highly range-restricted species induces unique challenges for species distribution modeling. In particular, bioclimatic covariates that are typically used to model species ranges at regional and continental scales may not show strong variation at scales of 100s and 10s of meters. This limits both the likelihood and usefulness of correlated occurrence to data typically used in distribution models. Here, we present analyses of species distributions, at 100 × 100 m resolution, for a highly range restricted salamander species (Shenandoah salamander, Plethodon shenandoah) and a closely related congener (red-backed salamander, Plethodon cinereus). We combined data across multiple survey types, account for seasonal variation in availability of our target species, and control for repeated surveys at locations- all typical challenges in range-scale monitoring datasets. We fit distribution models using generalized additive models that account for spatial covariates as well as unexplained spatial variation and spatial uncertainty. Our model accommodates different survey protocols using offsets and incorporates temporal variation in detection and availability resulting from survey-specific variation in temperature and precipitation. Our spatial random effect was crucial in identifying small-scale differences in the occurrence of each species and provides cell-specific estimates of uncertainty in the density of salamanders across the range. Counts of both species were seen to increase in the 3 days following a precipitation event. However, P. cinereus were observed even in extremely wet conditions, while surface activity of P. shenandoah was associated with a more narrow range. Our results demonstrate how a flexible analytical approach improves estimates of both distribution and uncertainty, and identify key abiotic relationships, even at small spatial scales and when scales of empirical data are mismatched. While our approach is especially valuable for species with small ranges, controlling for spatial autocorrelation, estimating spatial uncertainty, and incorporating survey-specific information in estimates can improve the reliability of distribution models in general.

Traversing the Great Lakes: Post‐glacial colonization by a widespread terrestrial salamander

AbstractAimsGlacial retreat at the end of the Pleistocene epoch opened vast expanses of emergent habitat in the northern hemisphere that were colonized by opportunistic taxa. However, species that undergo post‐glacial expansion may have originated from one or several glacial refugia. We inferred the post‐glacial expansion history of the Eastern Red‐backed Salamander (Plethodon cinereus), a fully terrestrial species with a vast distribution despite severe dispersal limitations. Previous studies indicated populations south of the glacial boundary at the eastern and western limits of the distribution were closely related, suggesting either multiple refugia or an extraordinary post‐glacial expansion event.LocationEastern North America.Taxon Plethodon cinereus (Green, 1818), Caudata: Plethodontidae.MethodsWe collected ddRAD‐seq data from 106 individuals throughout the distribution of P. cinereus. We estimated phylogeographic structure, including finer‐scale structure among the post‐glacial populations. To test the origins and routes of colonization, we used ecological niche modelling, population trees and analyses of directional range expansion.ResultsAnalyses supported our hypothesis of a southeastern glacial refugium, with northward expansion along the Eastern Seaboard prior to westward invasion into the Great Lakes region, including southwestern expansion into unglaciated areas at the western end of the distribution. However, a distinct subgroup in the northwestern portion of the range raises the possibility of a second refugium near the ice‐free Driftless Area.Main ConclusionsBased on our results, we hypothesize a southeastern refugium from which most of today's northern populations undertook extensive post‐glacial colonization. Our results indicate a geographically non‐linear colonization history for P. cinereus.

Population-level effects of prescribed fires on terrestrial salamanders

Prescribed fires are widely used for oak (Quercus) regeneration in hardwood forests, but the reported effects on terrestrial salamanders are variable, with some authors describing little to no effect of prescribed fire on salamanders and others reporting negative results. Factors such as variable fire intensity, sampling methodology, and imperfect salamander detection, which vary by study, may influence the effects of prescribed fires on salamanders. We examined the response of red-backed (Plethodon cinereus) and zig-zag (Plethodon dorsalis) salamanders to prescribed fire in forests of the Midwest U.S., accounting for changes in salamander behavior and detection by accounting for capture probability, and considering the influence of local fire intensity. We also explored the effects of fire on environmental variables (leaf litter depth, soil moisture, soil temperature, and soil pH) to identify possible mechanisms driving fire’s influence on salamanders. From spring 2019 through fall 2022, a total of 4922 salamander captures were recorded, representing 1939 individually marked salamanders at six coverboard grids in southern Indiana. Occurrence of fire affected survival and temporary emigration of salamanders, likely indicating that salamanders retreated underground, on two of the four grids. Fire occurrence and intensity did not significantly affect environmental variables except for leaf litter depth, which was negatively affected. However, soil moisture had a stronger influence on salamander capture probability than leaf litter. Overall, effects of prescribed fires on salamander populations in this study were inconsistent. In contrast, environmental factors had strong impacts on salamanders, but themselves exhibited either no relationship or a short-term relationship to prescribed fire variables. Notably, large salamander populations may be better able to tolerate prescribed fires. Such populations will be important to support during prescribed fire management to protect the species, especially as the effects of repeated prescribed fires and compounding effects are unknown. Understanding the interplay between forest management practices and wildlife habitat features such as soil moisture and leaf litter will be critical to identifying best practices for supporting wildlife populations reliant on such features during prescribed fires.

Specialist or refugee: Microhabitat use and competition between two sympatric woodland salamanders

AbstractCo‐occurrence between species may be mediated by ecological differences or competitive interactions, but the nature of these interactions can vary across spatial scales. At coarse scales, species may appear to broadly co‐occur, but at fine scales, particularly for species with small home ranges, species may utilize different aspects of the microhabitat such that co‐occurrence is limited and competitive interactions are reduced or eliminated. We investigated the microhabitat use of two morphologically and ecologically similar woodland salamanders (the Red‐backed Salamander, Plethodon cinereus, and the Northern Ravine Salamander, P. electromorphus) that are distributed throughout the Alleghany Plateau in eastern North America to determine whether they partition habitat at finer scales. We also tested if P. electromorphus is a specialist on habitat types that allow it to exclude P. cinereus, or if P. electromorphus uses lower quality habitat as a refuge to avoid P. cinereus. At a site of microsympatry, we repeatedly sampled plots during two seasons, each within a different area of the landscape, and used occupancy and abundance modeling to assess the extent of co‐occurrence and test whether the species use different microhabitat conditions. Plethodon cinereus greatly outnumbered P. electromorphus and occupied a greater proportion of plots, most plots where we detected P. electromorphus we also detected P. cinereus. In contrast to patterns documented in some montane species pairs, in which a high‐elevation habitat specialist excludes a low‐elevation generalist, we found that P. electromorphus primarily used microhabitat conditions favorable for both species, such as north‐facing slopes, yet did not have a negative effect on the abundances of P. cinereus. In the presence of an abundant competitor, microhabitat use of the less abundant species may narrow toward optimal conditions with sufficient resources to facilitate co‐occurrence.

Habitat selection and refuge-use by a color polymorphic salamander reveal behavioral niche differences.

Color polymorphic species provide an excellent opportunity to investigate the ecology and evolution of intraspecific niche differences. The red-backed salamander, Plethodon cinereus, is a fully terrestrial lungless salamander with two common color forms, striped and unstriped. Previous research suggests the morphs may be differentially adapted to surface and subsurface microhabitats, with the unstriped morph being more fossorial. This hypothesis predicts that the unstriped morph should be more sensitive to the risks of surface activity (e.g., thermal stress, dehydration, predation), and therefore be more selective than striped morphs when choosing soil surface microhabitats. To test this hypothesis, we experimentally manipulated leaf litter mass in small forest patches (~0.45 m2). Leaf litter addition reduced soil temperatures, buffered against changes in air temperature, and likely provided physical protection from predators. Over 3 years, we found that unstriped adults responded positively to leaf litter addition, but striped adults did not. In addition, unstriped morphs spent significantly more time in protective refuges (opaque, moistened tubes) than striped morphs in laboratory assays. Taken together, the field and laboratory results support the hypothesis that the unstriped morph is more sensitive to the risks of surface activity, and therefore is more likely to be fossorial. This difference in microhabitat use, combined with spatiotemporal variation in leaf litter accumulation on the forest floor, may play an important role in the maintenance of the polymorphism.

Influence of Natural Gas Pipeline Right-of-Ways on Eastern Red-Backed Salamander Occurrence in the Northern Appalachians

Abstract Road and pipeline infrastructure development for natural gas extraction often results in forest fragmentation, which could negatively influence habitat quality for many amphibian species. We investigated occurrence dynamics of the eastern red-backed salamander Plethodon cinereus in relation to natural gas pipeline rights-of-way (ROWs) and forest structure characteristics in northern Pennsylvania, USA. We sampled 80 sites across two study areas by using coverboards, with each site containing sampling plots at the center of the ROW, the edge of the ROW, and 10 m and 30 m into the adjacent forest. We assessed the influence of ROW age, ROW width, distance from ROW, and five forest structure characteristics on plot occupancy probability. Eastern red-backed salamander occupancy probability decreased with ROW age and increased with distance from ROW. Our results indicate that eastern red-backed salamanders are negatively influenced by forest fragmentation for natural gas ROWs. Moreover, responses were time dependent, with occupancy probability declining with ROW age. Due to low detections, we were unable to analyze data for the other amphibians and reptiles encountered during the study. Our capture data indicate that ROWs could improve habitat quality for some snake species, but additional research is needed to better understand the influence of ROWs on reptile species. To reduce future forest fragmentation and impacts on eastern red-backed salamander populations, managers could consider placing pipelines along existing linear clearings and enhancing the habitat quality of ROWs for salamanders.

Non-native earthworms increase the abundance and diet quality of a common woodland salamander in its northern range.

The online version contains supplementary material available at 10.1007/s10530-023-03168-3.

Projected northward shifts in eastern red-backed salamanders due to changing climate.

Many species' distributions are being impacted by the acceleration of climate change. Amphibians in particular serve numerous ecosystem functions and are useful indicators of environmental change. Understanding how their distributions have been impacted by climate change and will continue to be impacted is thus important to overall ecosystem health. Plethodon cinereus (Eastern Red-Backed Salamander) is a widespread species of lungless salamander (Plethodontidae) that ranges across northeastern North America. To better understand future potential lungless salamander range shifts, we quantify environmental favorability, the likelihood of membership in a set of sites where environmental conditions are favorable for a species, for P. cinereus in multiple time periods, and examine shifts in the species' distribution. First, utilizing a large data set of georeferenced records, we assessed which bioclimatic variables were associated with environmental favorability in P. cinereus. We then used species distribution modeling for two time periods (1961-1980 and 2001-2020) to determine whether there was a regional shift in environmental favorability in the past 60 years. Models were then used to project future distributions under eight climate change scenarios to quantify potential range shifts. Shifts were assessed using fuzzy logic, avoiding thresholds that oversimplify model predictions into artificial binary outputs. We found that P. cinereus presence is strongly associated with environmental stability. There has been a substantial northward shift in environmental favorability for P. cinereus between 1961-1980 and 2001-2020. This shift is predicted to continue by 2070, with larger shifts under higher greenhouse gas emission scenarios. As climate change accelerates, it is differentially impacting species but has especially strong impacts on dispersal-limited species. Our results show substantial northward shifts in climatic favorability in the last 60 years for P. cinereus, which are likely to be exacerbated by ongoing climate change. Since P. cinereus is dispersal-limited, these models may imply local extirpations along the southern modern range with limited northward dispersal. Continued monitoring of amphibians in the field will reveal microclimatic effects associated with climate change and the accuracy of the model predictions presented here.

Eastern red-backed salamanders (Plethodon cinereus) avoid chemical cues from snake predators regardless of colour morph or snake individual and size

Abstract Plethodontid salamanders often possess the ability to detect and avoid hazards in the environment which may be important for survival. We examined whether the two main colour morphs of Plethodon cinereus differ in their avoidance of cues from a potential predator (Thamnophis sirtalis) and whether P. cinereus avoided cues from T. sirtalis of different sizes (i.e., small vs. large). Plethodon cinereus avoided chemical cues deposited by T. sirtalis on a substrate; but avoidance did not differ between colour morphs of P. cinereus, snake sizes, or individual donor snakes.

Demography and space-use of Eastern Red-backed Salamanders (Plethodon cinereus) between mature and successional forests.

Space-use and demographic processes are critical to the persistence of populations across space and time. Despite their importance, estimates of these processes are often derived from a limited number of populations spanning broad habitat or environmental gradients. With increasing appreciation of the role fine-scale environmental variation in microgeographic adaptation, there is a need and value to assessing within-site variation in space-use and demographic patterns. In this study, we analyze 3 years of spatial capture-recapture data on the Eastern Red-backed Salamander collected from a mixed-use deciduous forest site in central Ohio, USA. Study plots were situated in both a mature forest stand and successional forest stand separated by <100-m distance. Our results showed that salamander density was reduced on successional plots, which corresponded with greater distance between nearest neighbors, less overlap in core use areas, greater space-use, and greater shifts in activity centers when compared to salamanders occupying the mature habitat. By contrast, individual growth rates of salamanders occupying the successional forest were significantly greater than salamanders in the mature forest. These estimates result in successional plot salamanders reaching maturity more than 1 year earlier than salamanders on the mature forest plots and increasing their estimated lifetime fecundity by as much as 43%. The patterns we observed in space-use and individual growth are likely the result of density-dependent processes, potentially reflecting differences in resource availability or quality. Our study highlights how fine-scale, within-site variation can shape population demographics. As research into the demographic and population consequences of climate change and habitat loss and alteration continue, future research should take care to acknowledge the role that fine-scale variation may play, especially for abiotically sensitive organisms with limited vagility.

Population genomic analyses support sympatric origins of parapatric morphs in a salamander.

In numerous clades, divergent sister species have largely non-overlapping geographic ranges. This pattern presumably arises because species diverged in allopatry or parapatry, prior to a subsequent contact. Here, we provide population-genomic evidence for the opposite scenario: previously sympatric ecotypes that have spatially separated into divergent monomorphic populations over large geographic scales (reverse sympatric scenario). We analyzed a North American salamander (Plethodon cinereus) with two color morphs that are broadly sympatric: striped (redback) and unstriped (leadback). Sympatric morphs can show considerable divergence in other traits, and many Plethodon species are fixed for a single morph. Long Island (New York) is unusual in having many pure redback and leadback populations that are spatially separated, with pure redback populations in the west and pure leadbacks in the east. Previous work showed that these pure-morph populations were genetically, morphologically, and ecologically divergent. Here, we performed a coalescent-based analysis of new data from 88,696 single-nucleotide polymorphisms to address the origins of these populations. This analysis strongly supports the monophyly of Long Island populations and their subsequent divergence into pure redback and pure leadback populations. Taken together, these results suggest that the formerly sympatric mainland morphs separated into parapatric populations on Long Island, reversing the conventional speciation scenario.

Evaluating Mechanisms of Short-term Woodland Salamander Response to Forest Management.

Contemporary forest management often requires meeting diverse ecological objectives including maintaining ecosystem function and promoting biodiversity through timber harvesting. Wildlife are essential in this process by providing ecological services that can facilitate forest resiliency in response to timber harvesting. However, the mechanisms driving species' responses remain ambiguous. The goal of this study was to assess mechanisms influencing eastern red-backed salamander (RBS; Plethodon cinereus) response to overstory cover removal. We evaluated two mitigation strategies for the RBS in response to overstory removal. We used a before-after-control-impact design to study how (1) retaining residual trees or (2) eliminating soil compaction affected RBS surface counts and body condition index (BCI) up to two-years post-treatment. Additionally, we assessed how surface counts of RBS were influenced by overstory tree cover. Surface counts of RBS were not strongly influenced by overstory removal when tree residuals were retained. Body condition index increased in treatments where harvest residuals were retained. In treatments where soil compaction was eliminated, surface counts and BCI were inversely related. Finally, surface counts from both mitigation strategies were not strongly influenced by overstory cover. Overall, both mitigation techniques appeared to ameliorate impacts of overstory removal on RBS. These results highlight the importance of understanding mechanisms driving species' responses to forest management. To reduce the perceived negative effects of overstory removal on RBS, incorporating these mitigation measures may contribute to the viability and stability of RBS populations. Incorporating species' life history traits into management strategies could increase continuity of ecological function and integrity through harvesting.

The Effects of Artificial Night Lighting on Tail Regeneration and Prey Consumption in a Nocturnal Salamander (Plethodon cinereus) and on the Behavior of Fruit Fly Prey (Drosophila virilis).

Simple SummaryAmphibians, including salamanders, are declining worldwide at an alarming rate due to a variety of factors that include habitat decline and destruction and environmental pollutants. Artificial light at night (ALAN) due to human activities is a nearly ubiquitous pollutant and can have serious consequences for amphibians. We examined the impact of ecologically-relevant levels of ALAN on tail regeneration in the eastern red-backed salamander, prey consumption by these salamanders and behavior of their fruit fly prey. We found that ALAN reduced the rate of salamander tail regeneration at some light levels above the naturally dark nocturnal illumination and increased the activity of their prey but not always in a simple, linear fashion. Thus, ALAN, even at very low levels, can influence the physiology and regeneration of a nocturnal salamander.As human development continues to encroach into natural habitats, artificial light at night (ALAN) has increasingly become a concern for wildlife. Nocturnal animals are especially vulnerable to ALAN, as the physiology and behavior of nocturnal species have evolved under conditions associated with predictably dark environments. Studies exposing amphibians to constant bright light provide evidence for changes to normal metabolism, growth, and behavior, but few of these studies have used treatments of dim ALAN comparable to that found in affected habitats. Eastern red-backed salamanders, Plethodon cinereus, use their tails for fat storage and communication, are capable of tail autotomy as an antipredator mechanism, and can regenerate the tail in its entirety. We examined the effect of different, ecologically-relevant intensities of ALAN on the rate of tail regeneration in adult P. cinereus. We hypothesized that ALAN would increase tail regeneration rates such that salamanders exposed to higher levels of light at night would regenerate tails faster than those exposed to lower light levels. In a controlled laboratory setting, we exposed salamanders (N = 76) in test chambers to nocturnal illuminations of 0.0001 lx (no ALAN, natural nocturnal illumination dark control), 0.01 lx (weak ALAN), 1 lx (moderate ALAN), or 100 lx (bright ALAN, equal to dim daytime and our day lighting treatment) for a period of 90 d immediately following tail autotomy. In addition, because these salamanders eat mostly live, moving prey, we investigated the impact of ALAN on the behavior of prey (Drosophila virilis) fed to the salamanders in our laboratory trials, which could alter feeding and regeneration rates in salamanders. We predicted that prey consumption would not be affected by ALAN and measured both prey consumption and prey behavior (activity) to examine the potential influence on regeneration. For tail regeneration, we found a non-monotonic response to ALAN, with salamanders exposed to nocturnal illuminations 0.1 lx and 100 lx regenerating tails significantly slower than salamanders in the 0.0001 lx or 1 lx treatments. Prey consumption did not differ among light treatments; however, fruit fly activity increased with increasing ALAN. These results suggest that ALAN influences regeneration rates, but the rate of regeneration is not dose-dependent and is not explained easily by prey consumption or movement of prey. We suggest that tail regeneration in these salamanders may involve a complex mechanism of altered gene expression and/or modulation of hormonal activity (corticosterone, melatonin, serotonin, and/or prolactin) at different intensities of nocturnal lighting.

Testing Assumptions in the Use of PIT Tags to Study Movement of Plethodon Salamanders

Studying the movements of organisms that live underground for at least a portion of their life history is challenging, given the state of current technology. Passive integrated transponders (PIT tags) provide a way to individually identify and, more recently, study the movement of smaller animals, including those that make subterranean movements. However, there are widespread assumptions of the use of PIT tags that remain problematic. We tested the effects of PIT-tag implantation on growth and survival, along with the effects of electromagnetic fields for reading PIT tags on behavior, of the smallest salamander that has been PIT-tagged: the Red-Backed Salamander. We found no effect of PIT tags on growth or survival. Using a mesocosm experiment, we also found that electromagnetic effects associated with reading PIT tags, had no effect on salamander behavior. Further, we describe a novel PIT antenna and soil mesocosm experimental arena for studying belowground movements of woodland salamanders. Collectively, these studies suggest that the use of PIT tags do not influence the growth, survival, or behavior of Red-Backed Salamanders. Given the challenges of studying salamanders that live underground and the impending changes in climate and landscapes, this research suggests that PIT tags remain a viable tool for studying the movement ecology of salamanders under global change.

Hybridization between the Woodland Salamanders Plethodon cinereus and P. electromorphus Is Not Widespread

A recent study reported widespread hybridization between the Eastern Red-backed Salamander (Plethodon cinereus) and the Northern Ravine Salamander (P. electromorphus) in northern Ohio. In this study, DNA sequence data were obtained from three nuclear loci and 20 single nucleotide polymorphisms (SNPs) were identified from the sequences. They found that 48 out of 90 individuals from 13 populations were hybrids, and in some localities every individual possessed an admixed genotype. As these results contradict our observations, and because levels of hybridization impact our interpretation of past and ongoing studies, we revisited the data. First we reanalyzed the original SNPs using STRUCTURE, then we repeated the analysis using haplotypes instead of SNPs. We found that K = 2 was best supported by both analyses, and they agree in recovering lower levels of hybridization than originally reported. For example, five populations in the original study identified as highly admixed or composed entirely of admixed genotypes we found to be pure P. cinereus or to lack evidence of extensive admixture. Similar results were obtained using NEWHYBRIDS and analyses based on gene trees. We conclude that while hybridization between P. cinereus and P. electromorphus occurs, it is much more restricted than originally reported.