Plethodon Cinereus Research Articles

Prescribed Fire Impacts to Amphibians and Reptiles in Shelterwood-harvested Oak-dominated Forests

As part of a larger study examining the role of prescribed fire in regenerating upland oaks (Quercus spp.), seasonal prescribed burns (winter, spring, summer, and unburned control) were applied to first-stage shelterwood-harvested stands on Horsepen Wildlife Management Area in the Virginia Piedmont in 1995. Because fire impacts are poorly documented for herpetofaunal communities, we surveyed these stands in 1996 capturing 133 individuals of ten species during over 12,720 pitfall trapnights. We found no significant differences in relative abundance of Eastern Red-backed Salamanders (Plethodon cinereus) (P = 0.26), American Toads (Bufo americanus (P = 0.93), or all amphibians combined (P = 0.25) among unburned shelterwood stands and those treated with winter, spring, or summer burns. Three species of reptiles (Northern Fence Lizard [Sceloporus undulatus], Ground Skink [Scincella lateralis], and Southeastern Five-lined Skink [Eumeces inexpectatus]) combined were captured more frequently in burned versus unburned stands (P = 0.02). Based on a stepwise multiple regression model, Eastern Red-backed Salamander captures were more strongly influenced by landscape variables (P = 0.0320), including distance to permanent water and mesic (i.e., eastern-northern) aspects, than by fire treatments (P = 0.26). Similar landscape models were not significant (P < 0.05) for toads or reptiles. Based on these results, prescribed fire may not be detrimental to herpetofaunal communities in oak dominated forests in the Virginia Piedmont.

Postautotomy tail movement differs between colour morphs of the red-backed salamander (Plethodon cinereus)

Striped and unstriped colour morphs of the eastern red-backed salamander,Plethodon cinereus, vary in their pre-attack behavioural response to predators, but it is unknown whether the morphs vary in post-attack strategies. Both morphs employ tail autotomy, a post-attack defensive mechanism enabling an individual to release a portion of their tail to facilitate escape from predation. Postautotomy tail movement diverts attention of a predator away from the individual’s body, so natural selection should favor vigorous tail movement in both colour morphs ofP. cinereus. We compared the degree of postautotomy tail movement between morphs following simulated predation. Striped individuals exhibited substantially longer and faster tail movement than unstriped individuals. Divergence in postautotomy tail movement may be a direct evolved response to variable predation pressure between colour morphs. Alternatively, tail movement may be constrained in the unstriped morph due to a genetic correlation with colouration (e.g., pleiotropy).

The relationship between climate and adult body size in redback salamanders (Plethodon cinereus)

Several biogeographic studies of salamanders have described relationships between salamander body size and climate. We specifically selected Plethodon cinereus as a widely distributed species that was well represented in museum collections to investigate the effects of warming climate on adult body size. We found a positive correlation between mean maximum July temperature and body size, and a negative correlation between precipitation of the driest month and body size. Surface‐collected adult P. cinereus were 2.3% larger in warmer counties on the coastal plain compared with cooler counties in the Appalachian Mountains. We compared salamanders collected between 1950 and 1970 versus those collected between 1980 and 2000 and found that body size increased 1.8% in counties on the coastal plain where mean maximum July temperatures had also increased by 0.5–1.2 °C. Warming temperatures alone, however, did not adequately account for the observed size increases, because body size also increased 1.3% in places that experienced less than 0.25 °C warming, but that difference was not statistically significant.

Climate-Mediated Competition in a High-Elevation Salamander Community

Abstract The distribution of the federally endangered Shenandoah Salamander (Plethodon shenandoah) is presumed to be limited by competition with the Red-backed Salamander (Plethodon cinereus). In particular, the current distribution of P. shenandoah is understood to be restricted to warmer and drier habitats because of interspecific interactions. These habitats may be particularly sensitive to climate change, though the influence of competition may also be affected by temperature and relative humidity. We investigated the response of P. shenandoah to competition with P. cinereus under four climate scenarios in 3-dimensional mesocosms. The results suggest that, although climate change may alleviate competitive pressure from P. cinereus, warmer temperatures may also significantly influence the persistence of the species across its known range.

Middle of the road: enhanced habitat for salamanders on unused logging roads

Context Amphibians are particularly susceptible to the effects of habitat loss and fragmentation. The construction and use of roads is among the most common sources of habitat fragmentation and can lead to serious population declines. Unused resource access roads, such as those formerly used for logging, can still negatively impact salamanders and reduce habitat quality through edge effects. Unfortunately, habitat rehabilitation and enhancement is rarely attempted on unused forest roads. Aims Our aim was to elaborate on a previous study that tested several types of woody debris to mitigate the negative impacts of forest roads by creating a novel habitat on an unused forest road in Algonquin Provincial Park. Here we focus solely on the use of large, squared timbers and their use by salamanders. Methods We tested the application of coarse woody debris (CWD) to the surface of an unused forest road. CWD were sampled for salamanders seven times during the 2011 field season. Local climatic variables were tested against salamander captures, and CWD size preferences and patterns of salamander aggregation under CWD were assessed. Key results We observed five salamander species and 415 individuals under timbers in the 2011 field season. Larger timbers (&gt;1m3) were preferred by all species observed and a significant proportion of animals were found in groups of two or more under larger timbers. High ambient temperature and low relative humidity negatively affected the number and species composition observed under timbers, suggesting that the efficiency of CWD as a survey method and enhanced habitat is season dependent. Implications Large timbers placed on unused forest roads may provide suitable refuges for migrating or dispersing forest salamanders while they attempt to cross the road. The tendency of salamanders to aggregate under CWD allows individual red efts to reduce water loss; however, red-backed salamanders are territorial and may drive off conspecifics. The use of large CWD may be an effective and low-cost method to rehabilitate unused forest roads and can be used to promote habitat connectivity for salamanders in targeted habitats, such as near wetlands, or for other species of concern.

Using Spatial Capture–Recapture to Elucidate Population Processes and Space-Use in Herpetological Studies

Abstract The cryptic behavior and ecology of herpetofauna make estimating the impacts of environmental change on demography difficult; yet, the ability to measure demographic relationships is essential for elucidating mechanisms leading to the population declines reported for herpetofauna worldwide. Recently developed spatial capture–recapture (SCR) methods are well suited to standard herpetofauna monitoring approaches. Individually identifying animals and their locations allows accurate estimates of population densities and survival. Spatial capture–recapture methods also allow estimation of parameters describing space-use and movement, which generally are expensive or difficult to obtain using other methods. In this paper, we discuss the basic components of SCR models, the available software for conducting analyses, and the experimental designs based on common herpetological survey methods. We then apply SCR models to Red-backed Salamander (Plethodon cinereus), to determine differences in density, survi...

Evaluating within‐population variability in behavior and demography for the adaptive potential of a dispersal‐limited species to climate change

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.

Assessing the impacts of the decline of Tsuga canadensis stands on two amphibian species in a New England forest

AbstractDisturbances such as outbreaks of herbivorous insects and pathogens can devastate unique habitats and directly reduce biodiversity. The foundation tree species Tsuga canadensis (eastern hemlock) is declining due to infestation by the nonnative insect Adelges tsugae (hemlock woolly adelgid). The decline and expected elimination of hemlock from northeastern U.S. forests is changing forest structure, function, and assemblages of associated species. We assessed 10 years of changes in occupancy, detection probability, and relative abundance of two species of terrestrial salamanders, Plethodon cinereus (eastern red‐back salamander) and Notopthalmus viridescens viridescens (eastern red‐spotted newt), to the experimental removal in 2005 of T. canadensis at Harvard Forest. Salamanders were sampled under cover boards and using visual encounter surveys before (2004) and after (2005, 2013, 2014) canopy manipulations in replicate 0.81‐ha plots. In 2004, occupancy of P. cinereus was 35% lower in stands dominated by T. canadensis than that in associated mixed‐hardwood control stands, whereas detection probability and estimated abundance of P. cinereus were, respectively, 60% and 100% greater in T. canadensis stands. Estimated abundance of N. v. viridescens in 2004 was 50% higher in T. canadensis stands. Removal of the T. canadensis canopy increased occupancy of P. cinereus but significantly reduced its estimated detection probability and abundance. Estimated abundance of N. v. viridescens also declined dramatically after canopy manipulations. Our results suggest that ten years after T. canadensis loss due to either the adelgid or pre‐emptive salvage logging, and 50–70 years later when these forests have become mid‐successional mixed deciduous stands, the abundance of these salamanders likely will be &lt;50% of their abundance in current, intact T. canadensis stands. This study adds to our understanding of how forest disturbance, directly and indirectly caused by invasive species, can contribute to declines in the relative abundance of amphibians.

Movement behaviour of woodland salamanders is repeatable and varies with forest age in a fragmented landscape

Landscape modification (loss, fragmentation and degradation of habitat) is a universal form of human-induced rapid environmental change that creates strong spatial variation in environmental conditions. Behaviourally mediated responses to landscape modification may generate behavioural divergence among populations along environmental gradients. We tested whether movement behaviour of red-backed salamanders, Plethodon cinereus, is repeatable and whether it varies within and between forest stands of different ages in a fragmented landscape in New York, U.S.A. Water loss is a major constraint on P. cinereus mobility, and forest fragmentation creates spatial variation in desiccation risk. We quantified multiple components of individual movement behaviour in an unfamiliar environment in the laboratory and used ordination to identify behavioural variation among individuals in multivariate space. We tested for repeatability of individual behaviour and then examined behavioural differences between individuals captured from populations at two spatial scales where desiccation risk was expected to vary: (1) between edge and interior locations within forest stands and (2) between forest stands varying in time since agricultural abandonment. Variation in movement behaviour among individuals was characterized by three components: total movement activity (e.g. surface activity, distance moved, boundary crossings), substrate use (time on moist soil versus dry sand) and latency to initial movement. Each behavioural component was significantly repeatable. At the population level, individuals from old forest stands (≥77 years) showed greater movement activity than individuals from young stands. Movement behaviour did not differ between individuals from edge and interior locations within forest stands. Our results demonstrate that P. cinereus individuals show consistent differences in movement behaviour in an unfamiliar environment. Moreover, we show that movement behaviour can be spatially structured within fragmented landscapes, but that behavioural divergence among populations depends on spatial scale.

Occupancy Rates and Detection Probabilities of Red-Backed Salamanders on the Virginia Fall Line

Occupancy Rates and Detection Probabilities of Red-Backed Salamanders on the Virginia Fall Line

Investigating temporal changes and effects of elevation on the prevalence of a rickettsial blood parasite in red-backed salamanders (Plethodon cinereus) in Virginia, USA

Amphibians are host to a number of vector-transmitted blood parasites. One parasite for which there is little information is a recently described intracellular bacteria in the order Rickettsiales, found in red-backed salamanders (Plethodon cinereus) in the eastern USA. This parasite was observed in 16.7 % of individuals across three states in the summer of 2008. Here, we resampled one of the locations from the 2008 study (Mountain Lake, Virginia, where site-level prevalence was 22 %) to determine if the incidence of infection has changed in 5 years. We also examined red-backed salamanders at seven other nearby sites that varied in elevation (670–1310 m) to ascertain if the pathogen shows any corresponding variation. We collected a total of 113 salamanders across the eight sites. We found no evidence that prevalence of the pathogen has changed; overall prevalence was 12.4 %, which was not significantly different from the 2008 estimate, and prevalence at the Mountain Lake site (28.6 %) also did not differ from the 2008 estimate. When comparing prevalence across elevations, we found that the highest-elevation sites tended to have highest levels of infections; one in five salamanders was infected with rickettsial bacteria at these locations. Ironically, these high-elevation sites appeared to have the highest salamander densities, and the largest salamanders, which usually point to healthy populations. Since salamander habitats in the Appalachian highlands are predicted to shrink in the future, our results suggest that the remnant, mountaintop populations will remain highly infected with this mysterious parasite.

COMPARING TWO TRANSECT METHODS FOR THE DETECTION OF RED-BACKED SALAMANDERS

Typical monitoring methods for terrestrial salamanders are subject to extensive variation, driven by the environmental conditions in effect during sampling. As rigorous salamander sampling methods are needed to monitor populations, there is a need to assess commonly used methods under a variety of environmental conditions. We hypothesized that of two methods used for capturing red-backed salamanders (Plethodon cinereus), leaf litter searches and natural cover searches, the latter would perform best for adults and in situations where moisture was limiting. We compared captures for paired transect surveys: one a leaf litter search and the other a natural cover search, relative to age, proximity to streams, rainfall events, vapor pressure deficit and season. We found that natural cover searches outperformed leaf litter searches when conducted away from streams and in the absence of rainfall. Natural cover searches performed better regardless of vapor pressure deficit and season (spring or fall). Natural cover searches detected more adults than juveniles. We recommend natural cover searches as more efficient than leaf litter searches.

Comparison of the Diets of Sympatric Erythristic and Striped Morphs ofPlethodon cinereus(Eastern Red-backed Salamander)

Abstract Plethodon cinereus (Eastern Red-backed Salamander) exhibits 3 distinct color morphs across eastern North America. The ecology of the least-common phenotype (i.e., erythristic morph) is largely unknown and no study has tested for ecological differences between sympatric erythristic and striped morphs. In this study, we compared dietary contents of striped and erythristic P. cinereus. We identified 553 prey items from 12 prey groups. Mean number and volume of prey per stomach did not differ between the 2 morphs. Important prey types were similar for both morphs and included Acari, formicids, Collembola, Diptera larvae, and adult Coleoptera. Additional studies regarding erythristic P. cinereus ecology will provide a better understanding of the role of this morph in natural populations.

Invasive Asian Earthworms Negatively Impact Keystone Terrestrial Salamanders.

Asian pheretimoid earthworms (e.g. Amynthas and Metaphire spp.) are invading North American forests and consuming the vital detrital layer that forest floor biota [including the keystone species Plethodon cinereus (Eastern Red-backed Salamander)], rely on for protection, food, and habitat. Plethodon cinereus population declines have been associated with leaf litter loss following the invasion of several exotic earthworm species, but there have been few studies on the specific interactions between pheretimoid earthworms and P. cinereus. Since some species of large and active pheretimoids spatially overlap with salamanders beneath natural cover objects and in detritus, they may distinctively compound the negative consequences of earthworm-mediated resource degradation by physically disturbing important salamander activities (foraging, mating, and egg brooding). We predicted that earthworms would exclude salamanders from high quality microhabitat, reduce foraging efficiency, and negatively affect salamander fitness. In laboratory trials, salamanders used lower quality microhabitat and consumed fewer flies in the presence of earthworms. In a natural field experiment, conducted on salamander populations from “non-invaded” and “pheretimoid invaded” sites in Ohio, salamanders and earthworms shared cover objects ~60% less than expected. Earthworm abundance was negatively associated with juvenile and male salamander abundance, but had no relationship with female salamander abundance. There was no effect of pheretimoid invasion on salamander body condition. Juvenile and non-resident male salamanders do not hold stable territories centered beneath cover objects such as rocks or logs, which results in reduced access to prey, greater risk of desiccation, and dispersal pressure. Habitat degradation and physical exclusion of salamanders from cover objects may hinder juvenile and male salamander performance, ultimately reducing recruitment and salamander abundance following Asian earthworm invasion.

Tongue and taste organ development in the ontogeny of direct-developing salamander Plethodon cinereus (Lissamphibia: Plethodontidae).

The latest research on direct developing caecilian and anuran species indicate presence of only one generation of taste organs during their ontogeny. This is distinct from indirect developing batrachians studied thus far, which possess taste buds in larvae and anatomically distinct taste discs in metamorphs. This study is a description of the tongue and taste organ morphology and development in direct developing salamander Plethodon cinereus (Plethodontidae) using histology and electron microscopy techniques. The results reveal two distinct stages tongue morphology (primary and secondary), similar to metamorphic urodeles, although only one stage of taste organ morphology. Taste disc sensory zones emerge on the surface of the oropharyngeal epithelium by the end of embryonic development, which coincides with maturation of the soft tongue. Taste organs occur in the epithelium of the tongue pad (where they are situated on the dermal papillae), the palate and the inner surface of the mandible and the maxilla. Plethodon cinereus embryos only possess taste disc type taste organs. Similar to the direct developing anuran Eleutherodactylus coqui (Eleutherodactylidae), these salamanders do not recapitulate larval taste bud morphology as an embryo. The lack of taste bud formation is probably a broadly distributed feature characteristic to direct developing batrachians. J. Morphol. 277:906-915, 2016. © 2016 Wiley Periodicals, Inc.

Can the eastern red‐backed salamander (Plethodon cinereus) persist in an acidified landscape?

AbstractHardwood forests of eastern North America have experienced decades of acidic deposition, leading to soil acidification where base cation supply was insufficient to neutralize acid inputs. Negative impacts of soil acidity on amphibians include disrupted embryonic development, lower growth rates, and habitat loss. However, some amphibians exhibit intraspecific variation in acid tolerance, suggesting the potential for local adaptation in areas where soils are naturally acidic. The eastern red‐backed salamander (Plethodon cinereus) is a highly abundant top predator of the northern hardwood forest floor. Early research found that P. cinereus was sensitive to acidic soils, avoiding substrates with pH &lt; 3.8 and experiencing decreased growth rates in acidic habitats. However, recent studies have documented P. cinereus populations in lower pH conditions than previously observed, suggesting some populations may persist in acidic conditions. Here, we evaluated relationships between organic horizon soil pH and P. cinereus abundance, adult health (body size and condition), and microhabitat selection, based on surveys of 34 hardwood forests in northeastern United States that encompass a regional soil pH gradient. We found no associations between soil pH and P. cinereus abundance or health, and observed that this salamander used substrates with pH similar to that available, suggesting that pH does not mediate their fine‐scale distributions. The strongest negative predictor of P. cinereus abundance was the presence of dusky salamanders (Desmognathus spp.), which were most abundant in the western Adirondacks. Our results indicate that P. cinereus occupies a wider range of soil pH than has been previously thought, which has implications for their functional role in forest food webs and nutrient cycles in acid‐impaired ecosystems. Tolerance of P. cinereus for more acidic habitats, including anthropogenically acidified forests, may be due to local adaptation in reproductively isolated populations and/or generalist life history traits that allow them to exploit a wider resource niche.

Silver-white Variants of the Eastern Red-backed Salamander, &lt;i&gt;Plethodon cinereus&lt;/i&gt;, from Eastern Canada

The Eastern Red-backed Salamander (Plethodon cinereus) is well known for its colour polymorphism, which includes 8 colour phenotypes: the red-backed (striped), lead-backed (unstriped), and erythristic morphs, as well as the iridistic, albino, leucistic, amelanistic, and melanistic anomalies. Among these, the prevalence and geographic distribution of rare morphs have received little attention. In this article, we report 2 occurrences of silver-white variants of the red-backed morph of P. cinereus from Québec, Canada. To our knowledge, these variations in stripe colour on the red-backed phenotype represent the first 2 documented mentions for eastern Canada.

Plethodon cinereus(Eastern Red-Backed Salamander) Not Affected by Long-Term Exposure to Soil Liming

Abstract The recovery of ecosystems affected by anthropogenic acidification is often a slow process, and one that is not always achievable through natural means. Application of carbonate materials to forest soils is being used more frequently to aid in the recovery of acidified ecosystems. However, few studies have addressed how the application of carbonate materials affects amphibians. We sampled field sites undergoing long-term application of high-calcium lime to investigate the effects of increases in soil pH on body condition and population demography of Plethodon cinereus (Eastern Red-backed Salamander). We found no effect of soil liming on body condition, population demographics, or density of surface-active Eastern Red-backed Salamanders. Our results are consistent with previous studies regarding the response of this species to soil liming, but unique in that they arise from an investigation of the long-term effects of liming exposure on density and demography in a wild population of Eastern Red-ba...

Responses of Eastern Red-backed Salamander (Plethodon cinereus) abundance 1 year after application of wood ash in a northern hardwood forest

Wood ash may be an effective soil amendment in North America to restore acidified and low-nutrient forest soils, but little research exists beyond its effects on soil and plants. Eastern Red-backed Salamander (Plethodon cinereus (Green, 1818)) abundance was assessed in a northern hardwood forest 1 year following an ash-addition field trial. Plots were established with fly ash and bottom ash treatments of 0, 1, 4, and 8 Mg·ha−1 (n = 4), and cover boards were positioned both with and without ash beneath. One year following ash additions, salamander abundance had increased under boards with fly ash beneath, and bottom ash had no effect. Soil pH and electrical conductivity increased under cover boards with ash beneath them and for uncovered soil, and the effects were strongest under cover boards with ash beneath. The effects of ash were generally stronger at higher dosages, and fly ash was stronger than bottom ash. The moisture holding capacity of fly ash was 60% higher than the soil and was 63% lower than the soil for bottom ash, but they had little effect on moisture of the forest floor. These results suggest that ash altered salamander abundance via soil pH and moisture and would not inhibit salamander movement over the forest floor.

Field-Testing a Standard Metabolic Rate Estimation Technique for Eastern Red-Backed Salamanders

Abstract Plethodontid salamanders are often monitored as indicator species for mature forest ecosystem health. In addition to relative abundance, differences in standard metabolic rate (SMR) between harvest treatments have been used to explain physiological stress in response to timber harvest. Nearly ubiquitous in forested stands throughout the northeastern United States, the Eastern Red-Backed Salamander (Plethodon cinereus) is often the focal species of such studies. In 2010, a predictive multiple regression equation was developed to calculate SMR of P. cinereus to 95% accuracy using salamander body mass and temperature. This method of SMR estimation has been implemented in field studies as a measure of salamander health. In these studies, temperature regime is the only variable measured, and SMR is calculated by standardizing on a 1-g salamander. In this study, we measured both body mass and temperature of each salamander encountered in harvested and unharvested stands and compared the published SMR c...