Northeastern Naturalist Vol. 26, No. 3 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 629 2019 NORTHEASTERN NATURALIST 26(3):629–640 A Survey of Terrestrially Active Salamanders from Two Different Woodlands at the Powdermill Nature Reserve, Pennsylvania Walter E. Meshaka Jr.1,*, Cordelia G. Lindsay2, Andrew L. Mack3, and Malcolm L. McCallum4 Abstract - Variation in habitat and microhabitat can affect salamander populations. We systematically surveyed terrestrially active salamanders in an old forest tract and a young forest tract at the Powdermill Nature Reserve in southwestern Pennsylvania during 2009– 2011. We examined the effect forest age had on the 2 salamander assemblage structures. While species compositions were similar, assemblage structures were uneven in both the old and young forest sites. Differential use of survey coverboards across species was detected at both sites. This difference in use may be best explained by ground-moisture levels as a function of coverboard distance to water. Counterintuitive findings of low numbers of Desmognathus ochrophaeus (Mountain Dusky Salamander) and Plethodon cinereus (Redbacked Salamander) compared to past studies at these 2 sites appears most likely related to differences in trapping methodology. Patterns of differential use of coverboards and inter-site differences in seasonal activity patterns point to P. glutinosus (Northern Slimy Salamander) as the least restricted species in terms of microhabitat conditions and most seasonally variable in activity. Introduction The Powdermill Nature Reserve (PNR) comprises protected forests, aquatic habitats, and grasslands in the northern Allegheny Mountains in Westmoreland County, PA (Meshaka et al. 2008). Westmoreland County is home to 17 species of salamanders (Meshaka and Collins 2012), and the PNR is home to 14 species of salamanders (Meshaka et al. 2008). The PNR can serve as a standard of representative habitats in a region long-affected by human activity, such as farming, strip mining, and logging. All such activities had taken place on the station before it was acquired for conservation purposes in 1956. We surveyed 2 forests of different ages to examine effects of ecological succession on salamander assemblage structure and composition. We compared our findings with those of a survey conducted in the early 1980s (Meshaka 2009) at sites near ours and contemporary collections made from sites on the PNR (Meshaka and Mortzfeldt 2014), which can be used to gauge responses by salamanders to type and age of habitat. Our expectation was 1Section of Zoology and Botany, State Museum of Pennsylvania, 300 North Street, Harrisburg, PA 17120. 2Carnegie Museum of Natural History, Powdermill Nature Reserve, 1795 Route 381, Rector, PA 15677. 3Pennsylvania State University-Altoona, 3000 Ivyside Park, Altoona, PA 16601. 4School of Agriculture and Applied Science, Langston University, Langston, OK 73050. *Corresponding author - wmeshaka@pa.gov. Manuscript Editor: Susan Herrick Northeastern Naturalist 630 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 Vol. 26, No. 3 dominance by Plethodon glutinosus (Green) (Northern Slimy Salamander) in the older hardwood forest, where it reaches its highest densities (Petranka 1998) Study Sites and Methods The study was conducted at the Powdermill Nature Reserve, an 890-ha field station, owned and operated by the Carnegie Museum of Natural History, located in Rector, Westmoreland County, PA. An older forested site, Calverly Lodge, and a young forested site, Upper Strip Mine (Fig. 1), typify mixed deciduous forest of Figure 1. Transects in the Powdermill Nature Reserve, Rector, Westmoreland County, PA, along (A) Calverly Lodge and (B) Upper Strip Mine. (C) A close-up of one of the coverboards used in the study. All images taken on 17 February 2012. Photographs © Cokie Lindsey. Northeastern Naturalist Vol. 26, No. 3 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 631 the northern Allegheny Mountains. The forest of Calverly Lodge, at an elevation of 463–475 m (Table 1), is composed of Quercus (oak)-dominated deciduous trees and Tsuga canadensis (L.) Carrière (Eastern Hemlock). An aerial photograph taken of the area in 1939 shows a dense forest with well-developed crowns. The Calverly Lodge site comprises a high-quality forest with respect to species diversity, height of trees, and total basal area of its trees. Areial photographs indicate that some of the Quercus rubra L. (Northern Red Oak) trees at this site likely exceed 100 years of age. The understory is vegetatively sparse with extensive rocks, leaf litter, and deadfall, and the forest floor is well-shaded in the summer. Powdermill Run, a creek, passes northwestward behind the lodge (Table 1). Immediately south of the creek, the forest floor is flat before inclining uphill southwest wardly. The Upper Strip Mine site is a younger forest and is located alongside Stripe Mine Road uphill and southeast from the bird-banding station (Table 1). This site is in recovery from strip mining. A seasonally dry stream is located downhill from and more or less parallel to the road. The site, at an elevation of 457–471 m (Table 1), is composed of a deciduous forest dominated by Acer (maple )and Liriodendron (tulip tree). An aerial photograph taken of the area in 1967 shows much of the area still bare since closure of the strip mine in 1947. Consequently, portions of the forest may be less than 40 years old. We screwed together six 1.0 m by 0.3 m sections of untreated hardwood 2.54 cm in thickness to form a single 1 m by 1 m coverboard that was 5.1 cm thick (Fig. 1). We placed 6 of those coverboards equidistantly at 10 m apart upon ground cleared of leaf litter to the mineral layer along a single transect at each of the 2 sites. In September 2008, we set coverboards on a transect perpendicular to Powdermill Run at Calverly Lodge and on a transect alongside and 10.7–16.8 m from the creek at Upper Strip Mine (Table 1). Coverboards were checked in the afternoon between the 1st and 29th of each month of April–October 2009–2011. All salamanders were captured by hand, identified to species, and released beside the coverboard. We recorded the presence of other vertebrates and miscellaneous behaviors. All statistical analyses were performed using MiniTab 13.3 (MiniTab, Inc. State College, PA). We compared the number of captures among the 3 sites using chi square (α = 0.05). Then, spatial relationships were analyzed. We tested data for normality using the Anderson–Darling normality test (α = 0.05). For any sets that violated normality assumptions, we transformed the data using the normality calculator function in MiniTab to allow use of parametric statistics before analysis using a general linear model (GLM) with a Tukey means comparison test. We reported these results, including the 95% confidence intervals (95% CI) for paired associations used in Tukey tests and the separate degrees of freedom from treatment and degrees of freedom from error (combined = total degrees of freedom). Confidence intervals that included zero were deemed not significantly different and those not including zero within their bounds were considered significantly different. For the GLM, we used α > 0.1 = reject, α < 0.05 = accept, and 0.1 > α > 0.05 = provisionally accepted. Northeastern Naturalist 632 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 Vol. 26, No. 3 Table 1. Physical parameters associated with coverboards at 2 sites at the Powdermill Nature Reserve, Rector, Westmoreland County, PA. The Esrignss average H RMS values represent the accuracy of the GPS reading (approximately less than 1 m). Esrignss Esrignss Esrignss Esrignss Site and Distance to Elevation latitude longitude Esrignss average average Esrignss board no. stream (m) (m) (°N) (°W) fixed date, time H RMS positions H Std. Dev. Calverly Lodge 01 28.27 463.23 40.146 -79.27 8/16/2018, 13:51 0.726735217 60 1.199734784 Calverly Lodge 02 42.00 463.26 40.146 -79.27 8/16/2018, 13:48 0.626546574 60 0.254931611 Calverly Lodge 03 52.05 464.16 40.146 -79.27 8/16/2018, 13:45 0.813208848 60 0.595194509 Calverly Lodge 04 62.28 465.57 40.146 -79.27 8/16/2018, 13:43 0.795426108 60 0.432735479 Calverly Lodge 05 73.12 470.40 40.146 -79.27 8/16/2018, 13:41 0.844236153 60 0.473161344 Calverly Lodge 06 84.37 475.04 40.146 -79.27 8/16/2018, 13:39 0.950215832 60 0.851214539 Strip Mine 01 11.59 471.54 40.159 -79.26 9/7/2018, 19:21 0.881981465 60 0.262834528 Strip Mine 02 10.72 469.89 40.160 -79.26 8/16/2018, 14:35 1.015152518 60 0.863505349 Strip Mine 03 12.26 468.75 40.160 -79.26 8/16/2018, 14:33 0.645858234 60 0.430890175 Strip Mine 04 15.29 467.23 40.160 -79.26 9/7/2018 ,19:32 0.753623249 60 0.648996458 Strip Mine 05 8.72 463.36 40.160 -79.26 8/16/2018, 14:31 0.758389953 60 0.522472861 Strip Mine 06 10.16 463.28 40.160 -79.26 8/16/2018, 14:30 0.689597638 60 0.777258083 Strip Mine 07 12.80 461.99 40.160 -79.26 9/7/2018, 20:09 0.544224560 60 0.198639247 Strip Mine 08 10.93 458.59 40.160 -79.26 8/16/2018, 14:28 0.712515281 60 0.722564062 Strip Mine 09 15.02 458.51 40.160 -79.26 8/16/2018, 14:26 0.769619710 60 0.255549774 Strip Mine 10 16.75 457.03 40.160 -79.26 8/16/2018, 14:25 0.700765427 60 0.190347924 Northeastern Naturalist Vol. 26, No. 3 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 633 Results Assemblage structures Desmognathus ochrophaeus Cope (Mountain Dusky Salamander), Plethodon cinereus (Green) (Red-backed Salamander), and Northern Slimy Salamander were present at both Calverly Lodge (27, 20, 55, respectively) and Upper Strip Mine (15, 35, 36, respectively). The distribution of captures among species was not even at Calverly Lodge (χ2 = 20.177, P < 0.001), where Northern Slimy Salamanders outnumbered Mountain Dusky Salamanders (χ2 = 22.56, P < 0.001) and Red-backed Salamanders (χ2 = 17.334, P = 0.003) (Fig. 2). Numbers of captures of Mountain Dusky Salamanders and Red-backed Salamanders were not significantly different from one another (χ2 = 1.042, P = 0.307). The distribution of captures differed significantly among species at Upper Strip Mine (χ2 = 9.79, P = 0.007), where the Mountain Dusky Salamander was encountered in numbers less than half that of each of the other 2 species (Fig. 2). To that end, the Mountain Dusky Salamander was encountered significantly less frequently than either the Northern Slimy Salamander (χ2 = 8.64, P = 0.003) or the Red-backed Salamander (χ2 = 8.00, P = 0.005). No significant difference was detected in the numbers of encounters of Northern Slimy Salamanders and Red-backed Salamanders (χ2 = 0.0014, P = 0.840). The encounter rate differed between sites, with only the Red-backed Salamander sharing similar Figure 2. Relative number of individuals of Desmognathus ochrophaeus (Mountain Dusky Salamander), Plethodon cinereus (Red-backed Salamander), and P. glutinosus (Northern Slimy Salamander) found under coverboards at 2 forested sites at the Powdermill Nature Reserve, Rector, Westmoreland County, PA, during 2009–2011. Northeastern Naturalist 634 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 Vol. 26, No. 3 encounter rates between sites (Fig. 2). Salamander relative abundance expressed per unit coverboard at Calverly Lodge was nearly twice that of Upper Strip Mine. Miscellaneous species Diadophis punctatus edwardsii (Merrem) (Northern Ringneck Snake) was found under cover at both Calverly Lodge (n = 1) and Upper Strip Mine (n = 2). We found a single Thamnophis sirtalis sirtalis (L.) (Eastern Garter Snake) under a coverboard at Strip Mine Road and a single Blarina sp. (shrew) under cover at Upper Strip Mine. Peromyscus leucopus (Rafinesque) (White-footed Mouse) was present at both sites; at Calverly Lodge, a family was present under cover on 2 occasions, a pair was present on 1 occasion, and a single individual was found on 1 occasion. Startled when uncovered, the pair of White-footed Mice bumped into the posterior ends of 2 juvenile Northern Slimy Salamanders, at which time both salamanders flipped their tails in an apparent attempt to hit the mice. At Upper Strip Mine, a family of White-footed Mice and a single shrew were encountered under coverboards. The Northern Slimy Salamander, an abundant member of this assemblage, is also a predator of smaller salamanders, including conspecifics. Opportunistic searches under natural cover in the immediate vicinity of the coverboards at both sites detected the presence of Plethodon wehrlei Fowler and Dunn (Wehrle’s Salamander) and Anaxyrus americanus (Holbrook) (American Toad) at Calverly Lodge. Spatial relationships Spatial distribution of salamanders along transects varied significantly at Calverly Lodge (A squared = 71.695, P < 0.001; Fig. 3). Abundance of salamanders varied by distance (F = 6.55, df = 5, dferror = 344, P < 0.001), month (F = 3.19, df = 6, dferror = 344, P = 0.005), and species (F = 11.32, df = 2, dferror = 344, P less than 0.001). Abundance varied among years but not quite statitstically significantly (F = 2.86, df = 2, dferror = 344, P = 0.059). Interaction between species and distance was correlated, creating multicollinearity; thus, it could not be included in the General Linear Model (GLM). A Tukey test did not reveal significant differences in abundance of the Mountain Dusky Salamander among the different coverboards; however, abundance dropped slightly (slope = -0.055) with distance (F = 2.90, P = 0.091). Occurrence of the Mountain Dusky Salamander was variable but not significantly different among coverboards (z = -1.73, P = 0.084). As distance increased from the creek, occurrence decreased but not significantly (slope = -0.0352, t = -1.76, P = 0.080). A Tukey test did not reveal significant differences in abundance of the Northern Slimy Salamander among different coverboards. Occurrence of the Northern Slimy Salamander did not vary among coverboards (z = -1.10, P = 0.271) nor with distance from the creek (t = 1.10, P = 0.273). Occurrence of the Red-backed Salamander, however, varied among coverboards (z = -2.13, P = 0.033) and it occurred slightly less frequently with distance (t = -2.24, P = 0.027, slope = -0.0394). The Red-backed Salamander became mildly less abundant as distance increased (t = 3.46, P = 0.001, slope = -0.0525). It was more common under coverboard 3 than all other coverboards except coverboard 1. The confidence interval for Northeastern Naturalist Vol. 26, No. 3 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 635 Figure 3. Numbers of Desmognathus ochrophaeus (Mountain Dusky Salamander), Plethodon cinereus (Red-backed Salamander), and P. glutinosus (Northern Slimy Salamander) found under each board at the (A) Calverly Lodge and (B) Upper Strip Mine at the Powdermill Nature Reserve, Rector, Westmoreland County, PA, during 2009–2011. Northeastern Naturalist 636 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 Vol. 26, No. 3 coverboard 1 overlapped with that of all other coverboards, hence no difference in occurrence existed between it and any other coverboard; however, its mean value was higher than all coverboards except coverboard 3. The spatial distribution of salamanders at Upper Strip Mine was not random along the transect (F = 4.34, df = 2, P = 0.013; Fig. 3). The GLM also detected significant differences among coverboards (F = 3.60, df = 9, P < 0.001) and months (F = 3.04, df = 6, P = 0.006) but not years (F = 0.560, df = 2, P = 0.573). Salamanders were more common in September than in July (Tukey 95% CI: -0.6350, -0.5720). All other months had similar abundance, and no significant differences were found among coverboards 1–6, 8, and 10. Inclusion of interaction variables was excluded because it resulted in multicollinearity. More salamanders were found under coverboard 7 than coverboard 1 (Tukey 95% CI: -0.6881, -0.0342), coverboard 5 (Tukey 95% CI: -0.6607, -0.0149), and coverboard 9 (Tukey 95% CI: 0.0469, 0.6854). No other significant differences were detected among coverboards. Non-normal distribution (A squared = 71.695, P < 0.001) of the Mountain Dusky Salamander distributions warranted transformation of the data. Subsequent analysis detected a non-random distribution of its use of coverboards (F = 2.06, dftreatment = 9, dferror = 199, P = 0.035); however, the Tukey tests did not identify significant differences. Individuals appeared to be most abundant under coverboard 2; however, the lower bound of the Tukey test was near the margin of significance (less than 0.03) for several of the distant coverboards (i.e., coverboards 9, 10), and a secondary mode of abundance of the Mountain Dusky Salamander was detected under coverboard 7. Abundance of the Northern Slimy Salamander did not differ significantly among coverboards (F = 1.79, dftreatment = 9, dferror = 199, P = 0.072). The Red-backed Salamander was not equally distributed among coverboards (F = 2.39, dftreatment = 9, dferror = 199, P = 0.014), whereby abundance under coverboard 8 was significantly greater than under coverboard 3 (Tukey 95% CI: -1.0710, -0.0032). Inclusion of interaction variables was excluded because it resulted in multicollinearity. Use of coverboards was in part related to the presence of other species. Presence of the Mountain Dusky Salamander interacted with the distance from the stream (board number) to inhibit occurrence of the Red-backed Salamander. However, the relationship was much tighter at Calverly Lodge (slope = -0.0194, r2 = 0.743, SEregression = 0.1913, P < 0.001), which runs perpendicular to Powdermill Run, than at Upper Strip Mine (slope = -0.0021; r2 = 0.046, SEregression = 0.341, P = 0.022), which runs nearly parallel to a stream. Coverboard use by the Northern Slimy Salamander occurred independently of either of the other species. Although the Northern Slimy Salamander was generally alone under cover at both sites, juveniles were often present under coverboards having more than one Northern Slimy Salamander. Seasonal activity Surface encounters of salamanders were distributed bimodally during the season at Calverly Lodge, Upper Strip Mine, and for both sites combined (Fig. 4). However, seasonal activity patterns varied among species and within species between Northeastern Naturalist Vol. 26, No. 3 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 637 Figure 4. Seasonal activity of of Desmognathus ochrophaeus (Mountain Dusky Salamander), Plethodon cinereus (Red-backed Salamander), and P. glutinosus (Northern Slimy Salamander) surveyed from coverboards at 2 forested sites, (A) Calverly Lodge and (B) Upper Strip Mine, and (C) the combined sample at the Powdermill Nature reserve, Rector, Westmoreland County, PA, during 2009–2011. Northeastern Naturalist 638 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 Vol. 26, No. 3 sites. The Mountain Dusky Salamander seemed to have been unimodal in its seasonal activity pattern, with strong pulses of surface activity in spring and early summer common to both sites. The Northern Slimy Salamander was most active during late summer through fall at Calverly Lodge and bimodally with a July drop in activity at Upper Strip Mine. Presumed courting pairs were evident in late summer and fall. At Calverly Lodge, an adult pair was found together under cover on 17 August 2011, and at Upper Strip Mine, a pair of adults was found under cover each on 21 September and 1 October 2011. The Red-backed Salamander was bimodal in its seasonal activity at both sites (Fig. 4). Discussion Our findings indicated that plethodons, especially the Northern Slimy Salamander, were the dominant component of the small assemblage of salamander at both sites. A systematic survey of amphibians was conducted in a young forest tract (alongside our Upper Strip Mine site) and the oldest forest tract in the PNR (north of our Calverly Lodge site) during 1982–1983 to assess the status of terrestrially active amphibian populations in various habitats of the PNR (Meshaka 2009). At that time, both sites were dominated by the Mountain Dusky Salamander, comprising more than 30% of assemblages of larger salamanders, and with very low numbers of the other species (Meshaka 2009). Proximity to Powdermill Run could explain the higher numbers of Mountain Dusky Salamanders and the detection of other streamside salamanders. Partially overlapping our study, Meshaka and Mortzfeldt (2014) systematically collected terrestrially active salamanders under cover at a different site on the PNR during 2008–2009. The flat, wet, Fagus (beech)–Acer (maple) forest site is partially surrounded by White Oak Creek. The forest has been intact for the past 80 years, excepting the creation of walking trails. Although different than either of our sites, Meshaka and Mortzfeldt’s (2014) site more closely resembled the Calverly Lodge site with respect to topography, moisture, and forest age. The assemblage found in that study was unevenly dominated by the Mountain Dusky Salamander, followed by Eurycea bislineata (Green) (Northern Two-lined Salamander), the Red-backed Salamander, and the Northern Slimy Salamander (Meshaka and Mortzfeldt 2014). Although very moist habitat is ideal for the Mountain Dusky Salamander, we do not know the extent to which differences in the 3 sampling techniques of these studies could result in differences in observed assemblage structures. Drift fences can be more productive for captures of certain salamanders than coverboards (Mills et al. 2013). Although native wood coverboards can match effectiveness of natural cover (Moore 2009), sampling results can vary as a result of coverboard design (Hesed 2012), such as direction (MacNeil and Williams 2012), age (Grasser and Smith 2014), and construction (MacNeil and Williams 2012), including thickness (Hesed 2012). To that end, the thickness of our coverboards would have provided thermal stability, and the size of the coverboards would have provided conditions that are warmer and drier than surrounding leaf litter (Hesed 2012). We do not know if the conditions associated with our coverboards affected the presence of the other Northeastern Naturalist Vol. 26, No. 3 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 639 2 species, both of which are also territorial (Petranka 1998), but they may have preferred ambient conditions that were different from those under our coverboards. Highest numbers of Red-backed Salamanders, and marginally so of Mountain Dusky Salamanders, under moist coverboards at Calverly Lodge corroborate this assertion. Likewise, the lead author made contemporary opportunistic collections of salamanders from under cover in the vicinity of Calverly Lodge and collections from the Calverly Lodge site after our study which yielded proportions of salamanders that were more similar to those of Meshaka’s (2009) study. Our coverboard design was also amenable to potential predators, especially small mammals and the Northern Slimy Salamander, a predator of small salamanders (Powders and Tietjen 1974) and conspecifics (Oliver 1967), at Calverly Lodge. For this reason, we leave open the possibility that the Mountain Dusky Salamander and the Red-backed Salamander, already compromised by the coverboard microhabitat, were also at risk of predation by the larger residential Northern Slimy Salamanders and small mammals. Activity pulses of these species are differentially affected by temperature and moisture (Petranka 1998) and thus, not surprisingly, can vary among sites or years. The general bimodality in seasonal activity often observed in the Northeast reflects the cool, wet spring and hot, dry summer followed by a cool, wet fall. Two previous studies on the PNR reflected this pattern generally. Meshaka (2009) found surface activity to be strongest in May and September. Elsewhere on the PNR, but more recently, pulses of surface activity were detected in May and August– September for the Mountain Dusky Salamander, July and September for the Northern Slimy Salamander, and May and October for the Red-backed Salamander (Meshaka and Mortzfledt 2014). Shifting modality in seasonal activity in the Mountain Dusky Salamander and the Northern Slimy Salamander could coincide with wetter summer conditions, whereas, mid-summer nesting might re-enforce bimodal seasonal activity in the Red-backed Salamander. The PNR salamander assemblage is species-rich even if highly uneven with respect to terrestrially moving species. Results of our systematic study were contrary to other assessments (Meshaka 2009, Meshaka and Mortzfeldt 2014), perhaps because of survey methodology and distance to water. However, inter-site differences in our study were evident in abundance and assemblage structure such that overall salamander numbers and those of the Northern Slimy Salamander specifically were higher in the older forest. Furthermore, coverboard choice underscored interspecific differences in microclimate requirements, the most broadly tolerant of which was the Northern Slimy Salamander, whose seasonal activity patterns were also the most variable among the species between sites. We are not surprised by the dominance of the Northern Slimy Salamander in the station’s oldest forest, Calverly Lodge, nor of the species we detected and their spatial distributions within the 2 sites. Biases are inherent to sampling techniques. In light of our interest in long-term demography of terrestrially active salamanders on the PNR, we advocate for duplicate coverboard studies using different designs concurrently to provide more than one kind of assessment of assemblage dynamics Northeastern Naturalist 640 W.E. Meshaka Jr., C.G. Lindsay, A.L. Mack, and M.L. McCallum 2019 Vol. 26, No. 3 as station forests continue to age in a northern Allegheny forest where intact, protected, and aged forests are rare. Acknowledgments We owe thanks to David Smith, past Director of the PNR, for his fostering a culture of research of which this project is a beneficiary. A vote of thanks is also due to Dr. John Wenzel, Director of the PNR, for active support in field research on the PNR and for his being a valued colleague and soundboard to research questions and ideas. We are grateful to James Whitacre for kindly mapping coverboard locations and Robert Leberman for his generosity in sharing his knowledge and insight of the Laurel Highlands. Literature Cited Grasser, C.N., and G.R. Smith. 2014. Effects of cover board age, season, and habitat on the observed abundance of Eastern Red-backed Salamander (Plethodon cinereus). Journal of North American Herpetology 1:53–58. Hesed, K.M. 2012. Uncovering salamander ecology: A review of coverboard designs. Journal of Herpetology 46:442–450. MacNeil, J.E., and R. N. Williams. 2012. Effectiveness of two artificial cover objects in sampling terrestrial salamanders. Herpetological Conservation and Biology 8:552–560. Meshaka, W.E., Jr. 2009. The terrestrial ecology of an Allegheny amphibian community: Implications for land management. The Maryland Naturalist 50:30–56. Meshaka, W.E., Jr., and J.T. Collins. 2012. A Pocket Guide to Pennsylvania Salamanders. 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Food habits of the White-throated Slimy Salamander in central Texas. Transactions of the Oklahoma Junior Academy of Science 1967:500–503. Petranka, J.W. 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington, DC. 587 pp. Powders, V.N., and W.L. Tietjen. 1974. The comparative food habits of sympatric and allopatric salamanders, Plethodon glutinosus and Plethodon jordani, in eastern Tennessee and adjacent areas. Herpetologica 30:167–175.