Plethodon Cinereus Research Articles

First report of Eastern Red-backed Salamander (<i>Plethodon cinereus</i>) on Newfoundland

The island of Newfoundland has no native amphibian taxa, although six species of Anura (i.e., frogs and toads) have been introduced since European colonisation, four of which have established self-sustaining populations. Here, we document Eastern Red-backed Salamander (Plethodon cinereus) on Newfoundland for the first time, in what appears to be a self-sustaining population near Conception Bay South. This is the first species of Caudata (i.e., newts and salamanders) to have been introduced to the island, as well as the first occurrence of Eastern Red-backed Salamander establishing a population outside its native range. The impact that this non-native species might have on forest ecosystems on Newfoundland is unclear and further study is required to determine whether eradication of the species from Newfoundland is necessary or feasible.

Computer vision for assessing species color pattern variation from web-based community science images

SummaryOpenly available community science digital vouchers provide a wealth of data to study phenotypic change across space and time. However, extracting phenotypic data from these resources requires significant human effort. Here, we demonstrate a workflow and computer vision model for automatically categorizing species color pattern from community science images. Our work is focused on documenting the striped/unstriped color polymorphism in the Eastern Red-backed Salamander (Plethodon cinereus). We used an ensemble convolutional neural network model to analyze this polymorphism in 20,318 iNaturalist images. Our model was highly accurate (∼98%) despite image heterogeneity. We used the resulting annotations to document extensive niche overlap between morphs, but wider niche breadth for striped morphs at the range-wide scale. Our work showcases key design principles for using machine learning with heterogeneous community science image data to address questions at an unprecedented scale.

Transposable Element Diversity Remains High in Gigantic Genomes

Transposable elements (TEs) are repetitive sequences of DNA that replicate and proliferate throughout genomes. Taken together, all the TEs in a genome form a diverse community of sequences, which can be studied to draw conclusions about genome evolution. TE diversity can be measured using models for ecological community diversity that consider species richness and evenness. Several models predict TE diversity decreasing as genomes expand because of selection against ectopic recombination and/or competition among TEs to garner host replicative machinery and evade host silencing mechanisms. Salamanders have some of the largest vertebrate genomes and highest TE loads. Salamanders of the genus Plethodon, in particular, have genomes that range in size from 20 to 70Gb. Here, we use Oxford Nanopore sequencing to generate low-coverage genomic sequences for four species of Plethodon that encompass two independent genome expansion events, one in the eastern clade (Plethodon cinereus, 29.3Gb vs. Plethodon glutinosus, 38.9Gb) and one in the western clade (Plethodon vehiculum, 46.4Gb vs Plethodon idahoensis, 67.0Gb). We classified the TEs in these genomes and found > 40 TE superfamilies, accounting for 22-27% of the genomes. We calculated Simpson's and Shannon's diversity indices to quantify overall TE diversity. In both pairwise comparisons, the diversity index values for the smaller and larger genome were almost identical. This result indicates that, when genomes reach extremely large sizes, they maintain high levels of TE diversity at the superfamily level, in contrast to predictions made by previous studies on smaller genomes.

Climate change and range restriction of common salamanders in eastern Canada and the United States

AbstractThe sensitivity of amphibian species to shifts in environmental conditions has been exhibited through long‐term population studies and the projection of ecological niche models under expected conditions. Species in biodiversity hotspots have been the focus of ample predictive modeling studies, while, despite their significant ecological value, wide‐ranging and common taxa have received less attention. We focused on predicting range restriction of the spotted salamander (Ambystoma maculatum), blue‐spotted salamander (A. laterale), four‐toed salamander (Hemidactylium scutatum), and red‐backed salamander (Plethodon cinereus) under future climate scenarios. Using bias‐corrected future climate data and biodiversity database records, we developed maximum entropy (MaxEnt) models under current conditions and for climate change projections in 2050 and 2070. We calculated positivity rates of species localities to represent proportions of habitat expected to remain climatically suitable with continued climate change. Models projected under future conditions predicted average positivity rates of 91% (89–93%) for the blue‐spotted salamander, 23% (2–41%) for the spotted salamander, 4% (0.7–9%) for the four‐toed salamander, and 61% (42–76%) for the red‐backed salamander. Range restriction increased with time and greenhouse gas concentration for the spotted salamander, four‐toed salamander, and red‐backed salamander. Common, widespread taxa that often receive less conservation resources than other species are at risk of experiencing significant losses to their climatic ranges as climate change continues. Efforts to maintain populations of species should be focused on regions expected to experience fewer climatic shifts such as the interior and northern zones of species' distributions.

Response of terrestrial salamanders to the decade following timber harvest in hardwood forests

Timber harvesting can have strong effects on terrestrial salamanders, which are critical components of forest ecosystems and indicators of environmental change. Effects of harvest methods such as clearcutting have been studied in the short term, but few studies have examined salamander trends throughout the decade following timber harvest. The effects of other harvest methods, such as patch cuts and shelterwoods, also remain unclear. We examined the effects of clearcuts, patch cuts, and shelterwood harvests on salamander relative abundance one year before and up to eleven years after harvest in clearcuts, shelterwoods, and patch cuts at the Hardwood Ecosystem Experiment (HEE). A total of 41,858 salamanders representing ten species were captured under artificial coverboards over this period with eastern red-back (Plethodon cinereus) and zigzag (Plethodon dorsalis) salamanders dominating. No significant declines occurred in the first three years following harvest, but salamander captures declined in the 4–6 and 7–11 year periods in clearcuts and patch cuts, and no signs of post-harvest recovery were found. Rapid vegetation recovery and high volumes of coarse woody debris (CWD) following harvest likely protected the microclimate used by salamanders; when CWD levels dropped 4–6 years after harvest, a corresponding decline in salamander captures occurred. These years also corresponded to a regional drought that likely increased temperature and decreased moisture in harvest openings, further impacting terrestrial salamanders. Neither the first nor second shelterwood harvests produced declines in salamanders, suggesting that canopy retention could prevent declines due to changing microclimate or drought. Our results highlight the need to consider compound effects of disturbances such as drought and canopy loss and the importance of long-term salamander monitoring following timber harvest to capture lagged effects and eventual recovery.

Does the thermal mismatch hypothesis predict disease outcomes in different morphs of a terrestrial salamander?

Many aspects of ectotherm physiology are temperature-dependent. The immune system of temperate-dwelling ectothermic host species is no exception and their immune function is often downregulated in cold temperatures. Likewise, species of ectothermic pathogens experience temperature-mediated effects on rates of transmission and/or virulence. Although seemingly straightforward, predicting the outcomes of ectothermic host-pathogen interactions is quite challenging. A recent hypothesis termed the thermal mismatch hypothesis posits that cool-adapted host species should be most susceptible to pathogen infection during warm temperature periods whereas warm-adapted host species should be most susceptible to pathogens during periods of cool temperatures. We explore this hypothesis using two ecologically and physiologically differentiated color morphs of the Eastern Red-backed Salamander (Plethodon cinereus) and a pathogenic chytrid fungus (Batrachochytrium dendrobatidis; hereafter "Bd") using a fully factorial laboratory experiment. At cool temperatures, unstriped salamanders (i.e., those that are tolerant of warm temperatures) had a significantly higher probability of Bd infection compared with cool-tolerant striped salamanders, consistent with the thermal mismatch hypothesis. However, we found no support for this hypothesis when salamanders were exposed to Bd at warm temperatures: the probability of Bd infection in the cool-tolerant striped salamanders was nearly identical in both cool and warm temperatures, opposite the predictions of the thermal mismatch hypothesis. Our results are most consistent with the fact that Bd grows poorly at warm temperatures. Alternatively, our data could indicate that the two color morphs do not differ in their tolerance to warm temperatures but that striped salamanders are more tolerant to cool temperatures than unstriped salamanders.

How does landscape permeability affect the movement of eastern red‐backed salamanders?

AbstractDemographic and genetic connectivity have important implications for population dynamics of wildlife and their ability to adapt to changing environments. Landscape features such as roads and streams can inhibit movement of species with poor dispersal capabilities and exacerbate the effects of habitat fragmentation. Terrestrial plethodontid salamanders are generally thought of as incapable of long‐distance movement and may vary in their motivation to move across a landscape by sex and size classes. To examine landscape permeability to movement, we assessed how distance and natural or man‐made landscape features affect recapture rates after displacement in red‐backed salamanders (Plethodon cinereus). We randomly assigned marked salamanders from 6 plots at Camp Karoondinha in Millmont, Pennsylvania, USA, to a control (placed under the same cover board) or a treatment (displaced 25 m or 50 m through the forest or across a stream or road). Of 290 marked individuals, we recaptured 29.8%. We recaptured more individuals in the control treatment compared to displaced individuals, with recapture rates decreasing with the presence of a road or stream. Landscape feature type and sex did not affect the probability of recapture, but developmental stage did, with adult salamanders more likely to be recaptured than juvenile salamanders. Our results suggest that landscape permeability is reduced for terrestrial salamanders when a landscape feature is present compared to open forest. Narrow, dirt roads present a similar challenge to a natural stream for red‐backed salamanders. Our study should be of value to land managers who are concerned about the negative effects of landscape permeability on terrestrial salamander populations.

Fecal Pellet Mass as a Function of Body Size, Color Morph, and Season in Plethodon cinereus (Green, 1818) (Amphibia: Plethodontidae)

In plethodontid salamanders, fecal pellets have a variety of behavioral functions related to chemical communication. We examined how fecal pellet mass in the Eastern Red-backed Salamander (Plethodon cinereus) varies with body size, color morph, and season. Fecal pellet mass increased with salamander size. Striped P. cinereus produced heavier fecal pellets than unstriped P. cinereus. Fecal pellet size tended to be greater in the spring than in the fall. Our results suggest further investigations of the factors governing fecal pellet size in plethodontid salamanders is warranted.

High landscape-level gene flow in the red-backed salamander (Plethodon cinereus)

Anthropogenic changes are expected to shape the genetic structure of many herpetofaunal populations. Indeed, genetic analyses are often applied in disturbed habitats because they can determine the loss of genetic diversity in isolated populations, identify barriers to dispersal and gene flow, and inform ways in which populations should be managed. As a result, there is a lack of baseline genetic data of organisms in relatively intact landscapes. Plethodontid salamanders are ideal for understanding fine-scale and landscape level genetic structure in continuous landscapes given their high abundance and their sensitivity to ecosystem changes. Given this, our study aimed to assess the landscape-level population structure in red-backed salamanders (Plethodon cinereus) within intact habitat. We investigated genetic variation in 479 individuals from three sites across roughly 26 km of intact hardwood forest. We tested for genetic structure among and within three sites using several metrics, including hierarchical Bayesian analysis, FST analysis, AMOVA, and isolation by genetic distance. The results revealed a single population throughout the study area and genetic structure within sites was not evident. Importantly, the sites in this study are part of a long-term project on the effects of standard silvicultural practices in the American Midwest, making it ideal for future re-evaluation of this red-backed salamander population. We now have baseline information to determine if any future disturbances or fragmentation may alter the genetic structure of this population, as well as how this population may change over time if the study area remains undisturbed.

Long-term occupancy dynamics of the threatened Cheat Mountain salamander and its competitors in relation to linear habitat fragmentation

Amphibians are declining globally and while many factors are contributing to this decline, habitat loss and degradation caused by climate and land use changes are among the most critical. Habitat degradation and increased interspecific competition are both concerns for long-term viability of the federally-threatened Cheat Mountain salamander (Plethodon nettingi) which is endemic to high elevations in West Virginia. In this study, we quantified the impacts of linear habitat fragmentation (i.e., a linear forest clearing for creation of a ski slope) on local colonization and extinction probabilities in adjacent forested habitat for the Cheat Mountain salamander and two co-occurring competitor species, eastern red-backed salamander (Plethodon cinereus) and Wehrle’s salamander (Plethodon wehrlei). We also quantified long-term changes in total occupancy of the species within the high elevation study area. We surveyed the salamander community annually from 1988 to 2021 using diurnal natural cover object searches at 43 plots, with 1988 representing three years following linear habitat fragmentation. For each species, we used dynamic occupancy models to identify and model influential covariates for initial occupancy, colonization, extinction, and detection probability. We found that distance to fragmentation was positively correlated with colonization probability for Cheat Mountain salamanders, indicating negative edge effects of the linear forest clearing. Distance to fragmentation was negatively correlated with colonization probability for eastern red-backed salamanders, potentially indicating this species benefited from increased solar radiation or reduced competition from Cheat Mountain salamanders. Predicted occupancy of eastern red-backed salamanders and Wehrle’s salamanders increased over the 34 year monitoring period, indicating potential for increased competitive interactions. Our study suggests that extensive linear habitat fragmentation could result in degraded habitat for Cheat Mountain salamanders in the adjacent forest, and that potential for interactions with competitor species is increasing in high elevation forest stands.

Rapid phenotypic change in a polymorphic salamander over 43\xa0years

Color polymorphic animals offer a unique system for studying intraspecific phenotypic responses to climate change. Discrete color morphs are easy to identify, and correlated trait responses of morphs can indicate how climate warming may facilitate long-term maintenance of polymorphisms. We use a historical dataset spanning 43 years to examine temporal shifts in color morph frequency and body size in response to climate in the Eastern Red-backed Salamander, Plethodon cinereus, which contains a widespread striped/unstriped color polymorphism. We created a pipeline to extract high-throughput trait data from fluid-preserved museum specimens where we batch-photographed salamanders, de-aggregated individual specimens from photographs, and solicited help of community scientists to score color morphs. We used a linear modeling framework that includes information about spatial population structure to demonstrate that color morph frequency and body size vary in response to climate, elevation, and over time, with an overall trend of higher frequency and decreased body size of the striped morph, but increased size of the unstriped morph. These surprising results suggest that morphs may be responding to multiple climate and geographic drivers through co-adapted morphological changes. This work highlights new practices of extracting trait data from museum specimens to demonstrate species phenotypes response to climate change.

Repeatability of water loss and gain rates of Plethodon cinereus

Abstract Water loss and gain rates of amphibians are important to understanding their ecology, especially for plethodontid salamanders. We report the first estimates of repeatability of relative water loss and gain rates of the two major colour morphs of the Eastern Red-backed Salamander, Plethodon cinereus, in the fall and spring seasons. Repeatability of relative water loss in P. cinereus was >0 in the fall but not in the spring. Repeatability of relative water gain was significant for all salamanders pooled in the fall, and was not repeatable in the spring. There were no apparent differences in repeatability of relative water loss or gain between the two colour morphs. Our results suggest that the repeatability of relative water loss and gain rates varies by season, but not by colour morph.

Are they arboreal? Climbing abilities and mechanics in the red-backed salamander (Plethodon cinereus).

While red-backed salamanders (Plethodon cinereus) are most often observed in terrestrial forested areas, several studies report arboreal substrate use and climbing behavior. However, salamanders do not have any of the anatomical features commonly observed in specialized climbing species (e.g., claws, setae, suction cups). Instead, salamanders cling to surfaces using the shear and adhesive properties of their mucous layer. In this study, we explore the capabilities and spatiotemporal gait patterns of arboreal locomotion in the red-backed salamander as they move across twelve substrate conditions ranging in diameter, orientation, and roughness. On arboreal substrates, red-backed salamanders decreased locomotor speed, stride frequency, phase and stride length, and increased duty factor and stride duration. Such responses have been observed in other non-salamander species and are posited to increase arboreal stability. Furthermore, these findings indicate that amphibian locomotion, or at least the locomotor behavior of the red-backed salamander, is not stereotyped and that some locomotor plasticity is possible in response to the demands of the external environment. However, red-backed salamanders were unable to locomote on any small-diameter or vertically-oriented coarse substrates. This finding provides strong evidence that the climbing abilities of red-backed salamanders are attributable solely to mucous adhesion and that this species is unable to produce the necessary external "gripping" forces to achieve fine-branch arboreal locomotion or scale substrates where adhesion is not possible. The red-backed salamander provides an ideal model for arboreal locomotor performance of anatomically-unspecialized amphibians and offers insight into transitionary stages in the evolution of arborealism in this lineage.

Beneath the Leaf-Litter: Can Salamander Personality Influence Forest-Floor Dynamics?

Ecological research demonstrates how intraspecific phenotypic variation can have consequences for community dynamics. However, the effects of animal personalities (i.e., intraspecific behavioral variation) on ecological processes remains relatively understudied. Using Red-backed Salamanders (Plethodon cinereus) and the detrital food web as a model system, we conducted a laboratory mesocosm experiment to explore whether or not the personality type of a top-level predator could affect the community structure within a complex, terrestrial food web. We used behavioral assays to investigate the repeatability of salamander behaviors, and classified individuals as either active or inactive. We then subjected laboratory mesocosms to one of four treatments for 3 mo: one active salamander, one inactive salamander, control (no salamander), and a pre-experimental reference. Our results indicate that the effect of P. cinereus on the detrital food web might be behaviorally mediated, with only the most active salamanders affecting community structure. Specifically, mesocosms housing active salamanders contained less diverse invertebrate communities than all other treatments. This difference was primarily driven by springtails, which were more abundant within mesocosms housing active salamanders. We also found that salamander personality was associated with cover board use inside mesocosms, with inactive salamanders exhibiting a more philopatric use of cover objects than active individuals. Leaf-litter degradation did not differ between treatments, indicating that the ecological effects of salamanders were too weak to influence basal resources within the detrital food web. Our study prompts further questions regarding the potential for animal personalities to influence ecological processes within terrestrial communities.

Experimental evaluation of spatial capture-recapture study design.

A principal challenge impeding strong inference in analyses of wild populations is the lack of robust and long-term data sets. Recent advancements in analytical tools used in wildlife science may increase our ability to integrate smaller data sets and enhance the statistical power of population estimates. One such advancement, the development of spatial capture-recapture (SCR) methods, explicitly accounts for differences in spatial study designs, making it possible to equate multiple study designs in one analysis. SCR has been shown to be robust to variation in design as long as minimal sampling guidance is adhered to. However, these expectations are based on simulation and have yet to be evaluated in wild populations. Here we conduct a rigorously designed field experiment by manipulating the arrangement of artificial cover objects (ACOs) used to collect data on red-backed salamanders (Plethodon cinereus) to empirically evaluate the effects of design configuration on inference made using SCR. Our results suggest that, using SCR, estimates of space use and detectability are sensitive to study design configuration, namely the spacing and extent of the array, and that caution is warranted when assigning biological interpretation to these parameters. However, estimates of population density remain robust to design except when the configuration of detectors grossly violates existing recommendations.

Assembly of the amphibian microbiome is influenced by the effects of land-use change on environmental reservoirs.

A growing focus in microbial ecology is understanding how beneficial microbiome function is created and maintained through various assembly mechanisms. This study explores the role of both the environment and disease in regulating the composition of microbial species in the soil and on amphibian hosts. We compared the microbial communities of Plethodon cinereus salamanders along a land-use gradient in the New York metropolitan area and paired these with associated soil cores. Additionally, we characterized the diversity of bacterial and fungal symbionts that putatively inhibit the pathogenic fungus Batrachochytrium dendrobatidis. We predicted that variation in skin microbial community composition would correlate with changes seen in the soil which functions as the regional species pool. We found that salamanders and soil share many microbial taxa but that these two communities exhibit differences in the relative abundances of the bacterial phyla Acidobacteria, Actinobacteria, and Proteobacteria and the fungal phyla Ascomycota and genus Basidiobolus. Microbial community composition varies with changes in land-use associated factors creating site-specific compositions. By employing a quantitative, null-based assembly model, we identified that dispersal limitation, variable selection, and drift guide assembly of microbes onto their skin, creating high dissimilarity between individuals with likely consequences in disease preventative function.

Repeatability of foraging behavior following a simulated predation attempt depends on color morph, sex, and foraging metric in Red-backed salamanders (Plethodon cinereus)

Behavioral repeatability greatly affects the capacity of an individual to respond to varying environments. When multiple behaviors within individuals are repeatable and correlated across time or across contexts, it is termed a behavioral syndrome. However, not all behaviors exhibit the same level of repeatability, and relatively few studies have examined repeatability in amphibians. We examined the repeatability of foraging behavior in the Eastern Red-backed salamander (Plethodon cinereus), a color-polymorphic terrestrial salamander, following a simulated predation attempt. We tested several hypotheses: (1) Simulated predation would negatively affect foraging, increasing latency to feed and decreasing the number of prey items eaten in a fixed time period compared to a control group; (2) Because striped color morphs of P. cinereus are more aggressive, striped individuals would exhibit “bold” behavior by resuming foraging sooner and consuming more prey; and (3) Foraging behavior would be more repeatable for males. We found that the predation treatment inhibited foraging behavior, although neither morphs nor sexes differed in either forging metric. The number of prey eaten was repeatable for all groups of salamanders. Latency to feed, however, was not repeatable for control salamanders. Simulated predation induced repeatable latencies, but when morphs and sexes were analyzed separately, only unstriped and male salamanders were repeatable, suggesting characteristics of these groups related to behavioral syndromes drive this response. We speculate that the greater repeatability of the unstriped morph’s latency to feed may result from more frequent encounters with predators in the leaf litter matrix while foraging. Striped salamanders from the source population, in turn, exhibit greater territorial success, and thus may experience more variation in encounters with predators and conspecifics over the course of their lifespans. Our results illustrate the need to carefully define the behavior and subset of the population to be tested when studying behavioral repeatability or behavioral syndromes.

Infection with Batrachochytrium dendrobatidis reduces salamander capacity to mount a cell-mediated immune response.

The vertebrate immune system is a costly defense system that is responsible for preventing and eliminating parasites and pathogens. Theory predicts that hosts experience tradeoffs associated with immune deployment and other physiological functions. Although empirical evidence for immune-physiology tradeoffs are well documented in the literature, fewer studies have examined tradeoffs within the immune system in wild vertebrates. We explored the topic of concomitant immune challenges in amphibians by exposing salamanders (Plethodon cinereus) to a fungal pathogen Batrachochytrium dendrobatidis (hereafter "Bd") and then to phytohemagglutinin (hereafter "PHA"). We measured Bd infection using quantitative PCR and used measurements of the tail thickness at the PHA injection site as an estimate of skin swelling. We tested whether Bd reduced the salamander's capacity to mount an immune response towards PHA or whether Bd would stimulate immune activity and thereby increase the response towards PHA. Salamanders that were infected with Bd had a reduced skin-swelling when injected with PHA compared to noninfected salamanders, a result that is consistent with the hypothesis that Bd-infected salamanders have lower immunocompetence than noninfected salamanders. We also found that PHA-induced swelling response was negatively associated with Bd infection abundance (i.e., the infection burden of all exposed salamanders, including those that were exposed but not infected), indicating that salamanders with a higher infection abundance had the lowest swelling response to PHA. Our results suggest that individuals of P. cinereus might experience an energetic tradeoff between successfully fighting off Bd and mounting an immune response towards PHA.

Salamander survey of Jakey Hollow Natural Area, Columbia County, Pennsylvania

AbstractWith current declines in amphibian and reptile populations, herpetological surveys of protected natural areas are crucial to management of these organisms, especially as natural areas become more insular due to surrounding land use. Jakey Hollow Natural Area (JHNA) is a locally significant biological site in Columbia County, PA, consisting of wooded slopes bisected by a small-order stream. This report presents the results of a salamander survey at JHNA during the spring of 2020. Cover object surveys revealed a total of 190 salamanders belonging to seven species. The most common salamanders observed at JHNA were Plethodon cinereus and Desmognathus ochrophaeus, which typify other surveys of similar habitats in Pennsylvania. Other species found in lower abundance were D. fuscus, Eurycea bislineata, Gyrinophilus porphyriticus, P. glutinosus, and Pseudotriton ruber. Future surveys utilizing other techniques may reveal the presence of additional salamander species at this site.

Maternal Body Size and Condition Predict Measures of Reproductive Success and Future Reproductive Allocation in Territorial Eastern Red-Backed Salamanders

Female fitness is often influenced by trade-offs in energy allocated to reproduction or self-maintenance that is dependent upon the life history of the organism. Maternal body size and condition are factors that are often positively correlated with clutch size and offspring size. We investigated whether maternal body size (snout–vent length, SVL) and condition (residual of mass on SVL) could predict measures of reproductive success including clutch size, offspring body size (SVL), and offspring survival in the territorial Eastern Red-backed Salamander (Plethodon cinereus). Brooding females and their eggs were collected and then housed in a controlled laboratory setting. We recorded the number of eggs found with each female and separated offspring from females once hatched. At 40 d after hatching, we measured the SVL and mass of females and juveniles. Maternal SVL, but not condition, was positively related to clutch size and offspring SVL; however, there was no significant relationship between clutch size and offspring SVL, indicating no trade-off between number and body size of offspring. In the laboratory, survival of the offspring to 185 d post-hatching was positively related to maternal SVL and negatively related to maternal condition. Also, the mean SVL, measured on day 40, of the offspring that survived to 185 d was significantly greater than the SVL of the offspring that did not survive, indicating a benefit of larger offspring body size. Finally, subsequent egg production (number of new oocytes produced by females following the current reproductive bout) was positively related to maternal SVL and condition (measured 185 d after hatching their previous clutches). These results suggest that larger maternal body size and condition of Eastern Red-backed Salamanders provide increased reproductive success through the number and body size of offspring produced in current or subsequent reproductive bouts and in offspring survival. Large body size in territorial animals like P. cinereus provides benefits for holding high quality territories as well as for increased reproductive success.