United States

Department of 

Agriculture

Forest Service

Northern 

Research Station

Research Paper NRS-10

Reptile, Amphibian, and Small 

Mammal Species Associated 

with Natural Gas Development 

in the Monongahela National 

Forest, West Virginia

Kurtis R. Moseley

W. Mark Ford

John W. Edwards

Mary B. Adams
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Abstract

Burgeoning energy demand in the United States has led to increased natural gas 

exploration in the Appalachian Basin. Despite increasing natural gas development in the 

region, data about its impacts to wildlife are lacking. Our objective was to assess past and 

ongoing natural gas development impacts on reptiles, amphibians, and small mammals 

in the Monongahela National Forest in West Virginia. We sampled 40 gas well sites and 

compared amphibian, reptile, and small mammal captures among active producing, plugged 

(inactive), and storage well types. Total species richness and diversity were greater at 

storage gas well sites than at plugged wells. Although natural gas development adversely 

impacts moisture-sensitive woodland salamanders, our results suggest that maintained gas 

well openings may benefi t other herpetofauna and small mammal species that use early 

successional habitat within predominately forested central Appalachian landscapes.

Cover Photo

Gas well on the Fernow Experimental Forest. Photo by the U.S. Forest Service.

Manuscript received for publication 12 August 2009
1

INTRODUCTION

Burgeoning energy demand and concomitant interest in reducing 

dependence on foreign fossil fuels have prompted increased exploration 

and production eff orts within the United States (U.S. Department 

of Energy 2003, 2005). Th e Appalachian Basin, centered along the 

Appalachian Mountains from New York to Alabama, is receiving 

increasing attention as an economically recoverable natural gas source 

with an estimated 2 tcm of undeveloped natural gas resources within 

the basin (Milici et al. 2003). Over the past decade, advances in 

drilling technology, increasing natural gas prices, existing pipeline 

infrastructure, and proximity to natural gas markets in the heavily 

urbanized northeastern United States have prompted increased 

exploration and subsequent production in the Appalachian Basin. 

Consequently, natural gas development in the Appalachian Basin is 

projected to continue to increase, particularly in the central Appalachian 

region of Ohio, Pennsylvania, Virginia, and West Virginia (U.S. 

Department of Energy 2005). Despite the current and projected 

growth of natural gas development within this region, many of its 

environmental impacts, particularly on wildlife species, largely are 

undocumented (Energy and Biodiversity Initiative 2007). Because 

natural gas development involves a complete clearing of small areas 

of forest for wellhead installation and associated pipeline and road 

development, wildlife sensitive to local-scale habitat alterations, such as 

herpetofauna (deMaynadier and Hunter 1995, Russell et al. 2004) and 

small mammals (Kaminski et al. 2007, Kirkland 1977), may be aff ected.

Forest openings associated with natural gas development undoubtedly 

increase edge and early successional habitats within the predominately 

forested central Appalachian landscape. Forest edge, defi ned as a 

transition zone, abrupt or gradual, between two adjacent ecosystems or 

vegetative communities (Murcia 1995), and early successional habitat 

creation and maintenance often enhance diversity of small mammals 

(Johnson et al. 1979, Lowell and Geis 1983, Menzel et al. 1999) 

and reptiles (Greenberg 2001, Kjoss and Litvaitis 2001, Pais et al. 

1988, Ross et al. 2000). For example, edge habitat between northern 

hardwood forests and wildlife openings in the southern Appalachians 

of North Carolina supported greater small mammal diversity than did 

forest interior habitat (Menzel et al. 1999). Similarly, species richness 

and diversity of small mammals were greater in powerline rights-ofway and edge than adjacent oak (Quercus spp.)-hickory (Carya spp.) 

forest in eastern Tennessee (Johnson et al. 1979). Although edge and 

early successional habitat use by reptile assemblages in the central 

Appalachian region has received scant attention, reptile diversity and 

richness often increase following forest canopy removal (Greenberg 

2001, Ross et al. 2000).

Th e Authors

KURTIS R. MOSELEY was a graduate 

research assistant, Division of Forestry 

and Natural Resources, West Virginia 

University, Morgantown, WV, and 

currently is wildlife biologist for the United 

States Marine Corps, Quantico, VA.

W. MARK FORD is a research wildlife 

biologist, formerly with the U.S. Forest 

Service, Northern Research Station, 

currently with the U.S. Army Engineer 

Research and Development Center, 

Vicksburg, MS.

JOHN W. EDWARDS is an associate 

professor, Division of Forestry and Natural 

Resources, West Virginia University, 

Morgantown, WV.

MARY B. ADAMS is a supervisory soil 

scientist, U.S. Forest Service, Northern 

Research Station, Parsons, WV.
2

Alternatively, gas well openings can adversely impact 

species dependent on moist, cool microclimate 

conditions, such as woodland salamanders, characteristic 

of continuous woody canopy cover (deMaynadier and 

Hunter 1995, Russell et al. 2004). In the Appalachian 

Mountains, woodland salamanders can exhibit drastic 

population declines after forest canopy removal following 

disturbances such as timber harvesting (Ash 1997, Ford 

et al. 2002, Petranka et al. 1993, Ross et al. 2000); 

subsequent salamander population recovery closely 

parallels reestablishment of important microhabitats, 

such as coarse woody debris (CWD) and leaf litter, as 

the forest regenerates (deMaynadier and Hunter 1995, 

Pough et al. 1987). However, timber harvest represents 

a more ephemeral form of forest disturbance than 

openings associated with natural gas development. Gas 

well openings are maintained in an early successional 

state through regular mowing for prolonged periods 

(i.e., the production life of the well). Such openings, in 

addition to associated roads for other similar types of 

disturbance and conversion, have been hypothesized to 

hinder salamander dispersal (Rittenhouse and Semlitsch 

2006, Rothermel and Semlitsch 2002), possibly resulting 

in long-term declines of local populations (Cushman 

2006).

Small mammals (Carey and Harrington 2001, Pearce 

and Venier 2005) and amphibians (Welsh and Ollivier 

1998, Welsh and Droege 2001) often serve as indicators 

of sustainable forest management. Th erefore, we need 

to understand how habitat alterations associated with 

natural gas development within predominately forested 

landscapes infl uence abundance and diversity of these 

taxonomic groups so that natural resource managers 

can more eff ectively mitigate natural gas development 

impacts in the central Appalachians and elsewhere. 

Accordingly, our goal was to conduct a survey of 

amphibians, reptiles, and small mammals associated 

with natural gas development in the Monongahela 

National Forest (MNF) in West Virginia. Additionally, 

we examined (1) diff erences in amphibian and reptile 

captures between gas well openings and adjacent forest 

habitat, (2) diff erences in small mammal captures 

among gas well openings, gas well opening/forest edge, 

and adjacent forest habitat, and (3) infl uence of gas 

well site status and aspect on sampled taxa and species 

diversity and richness. Because amphibians require 

moist, cool microhabitats (Duellman and Trueb 1994), 

we hypothesized that amphibian capture rate would 

be greater in adjacent forested habitat than in gas well 

openings. Conversely, we predicted that reptile capture 

rate would be greater in gas well openings because 

increased solar radiation would allow increased basking 

opportunity (Greenberg 2001, Seigel et al. 1987). For 

small mammals, we predicted that total captures would 

not diff er among gas well openings, gas well opening/

forest edge, and adjacent forest because, as a taxonomic 

group, small mammals exhibit broad habitat selection 

in the central Appalachian Mountain region (Kirkland 

1977, 1990, Lidicker 1999). For the infl uence of 

gas well site status, we hypothesized that reptile and 

amphibian capture rate, small mammal captures, and 

species diversity and richness would be greater at plugged 

well sites because these sites do not experience repeated 

mechanized disturbance, therefore providing increased 

microhabitat.

STUDY SITE

Th e 364,225-ha MNF is located within the central 

Appalachian Mountains of West Virginia in portions 

of 10 counties. Most of the MNF is located in the 

Allegheny Mountains and Plateau physiographic 

subprovince, where forests at lower to mid-elevations 

are dominated by the mixed mesophytic hardwood 

type consisting of sugar maple (Acer saccharum), red 

maple (A. rubrum), northern red oak (Quercus rubra), 

chestnut oak (Q. prinus), yellow-poplar (Liriodendron 

tulipifera), American beech (Fagus grandifolia), sweet 

birch (Betula lenta), black cherry (Prunus serotina), and 

basswood (Tilia americana; Madarish et al. 2002). At 

approximately 900 to 1,100 m elevation and depending 

upon aspect and landform position, the forest transitions 

to northern hardwood or northern hardwood—montane 

boreal assemblages of sugar maple, American beech, 

yellow birch (B. alleghaniensis), eastern hemlock (Tsuga 

canadensis), and red spruce (Picea rubens; Stephenson 

1993). Annual precipitation is approximately 152 cm, 

much of which occurs as snow in the winter. Th e frostfree period ranges from 90 to 150 days depending on 

elevation (Stephenson 1993).
3

Approximately 93 percent of the MNF is forested, 

but it also contains small nonforested areas limited to 

small livestock grazing allotments, wildlife food plots or 

maintained grassy openings, and natural gas well sites 

(U.S. Forest Service 2006). Currently, there are 96 gas 

wells on the MNF consisting of actively producing, 

plugged (inactive), and storage sites. Storage sites are 

natural underground cavities where methane gas is 

maintained in reserve during periods of low demand, 

such as the summer. Gas well sites were established 

between 1957 and 2001 with the majority established 

before 1970. Since establishment, active producing and 

storage gas well sites have been maintained as grass- and 

forb-dominated openings by biennial mowing. Gas well 

opening area on sampled sites ranged from 0.15 to 1.44 

ha and elevation ranged from 648 to 1,185 m. Gas well 

opening distance from streams ranged from 30 to 645 m. 

All gas well openings are connected by gas well pipeline 

rights-of-way or gravel access roads.

METHODS

We used ArcMAP 9.0 (ESRI, Redlands, CA) to 

randomly select a subsample of the 96 natural gas 

well sites on the MNF representing equal numbers of 

the three main well types: plugged, producing, and 

storage wells. However, due to logistical constraints, 

we could not survey some selected sites. Overall, we 

sampled 11 plugged, 13 producing, and 16 storage 

natural gas well sites. We conducted amphibian, reptile, 

and small mammal surveys at all 40 natural gas well 

sites throughout the MNF from June to August 2006 

(Fig. 1). We sampled reptiles and amphibians along 

four 1-m-wide strip transects originating from the 

center of each gas well and extending 100 m into the 

forest interior from the gas well edge. Each transect was 

oriented toward one of the four cardinal directions. 

Some transect locations deviated from cardinal directions 

(maximum 45°) to maintain a distance of at least 50 m 

Figure 1.—Plugged (inactive; n = 11), storage (n = 16), and producing (n = 13) gas well sites sampled 

for reptiles, amphibians, and small mammals within the Monongahela National Forest in West Virginia, 

June-August 2006.
4

from connected linear openings of roads and gas lines. 

We overturned and searched all cover items encountered 

along each transect for reptile and amphibian presence 

between 0800 and 1800 hr. We conducted searches of 

each transect only once during our study and did not 

conduct searches during precipitation events. Th erefore, 

our salamander capture estimates are conservative relative 

to similar studies in the region that sampled during 

cooler, moister weather conditions. We did not search 

leaf litter due to time constraints. However, amphibian 

captures under natural cover objects have been correlated 

with independent estimates of abundance (DeGraaf 

and Yamasaki 1992, Smith and Petranka 2000). When 

we encountered an animal, we identifi ed it to species 

and recorded the location of capture as forest or gas 

well opening habitat. Precipitation during our sampling 

period totaled 55.1 cm (20-yr average = 40.4 cm).

To sample small mammal populations at gas well sites, 

we established three trap lines: one at the center, one at 

the edge of gas well openings, and one 50 m from the gas 

well edge within adjacent mixed mesophytic, northern 

hardwood, or red spruce forest (Menzel et al. 1999). 

Trap lines consisted of 20 snap traps separated by 1 m 

and baited with peanut butter. We opened traps for one 

3-day period at each gas well site, checking traps daily 

and reapplying bait as necessary. We recorded species for 

each individual captured. In the Northeast and midAtlantic region, deer mice (Peromyscus maniculatus) and 

white-footed mice (P. leucopus) are diffi cult to distinguish 

based on pelage and external morphology (Aquadro and 

Patton 1980, Laerm and Boone 1994). Th erefore, we 

grouped these species as Peromyscus spp. for analyses. We 

conducted trapping and collection under West Virginia 

Department of Natural Resources scientifi c research 

permit 2006.036 and West Virginia University ACUC 

number 06-0506.

We used ArcMAP 9.0 to determine elevation, aspect, 

road density, gas well opening area (m2

), and gas well 

site distance to nearest stream (m; any intermittent 

or perennial lotic waterbody identifi ed on the USGS 

National Hydrography Dataset http://nhd.usgs.gov/). 

U.S. Forest Service personnel recorded gas well site 

locations using a global positioning system. We derived 

values for elevation and aspect using a 30 × 30 m digital 

elevation model (West Virginia GIS technical center, 

West Virginia University). We calculated road density 

as total road length (m) within 100-m buff ers around 

sample sites. We calculated combined amphibian, 

reptile, and small mammal species richness and 

Shannon’s Diversity Index (H’) for each gas well site. We 

categorized gas well sites as mesic, xeric, and intermediate 

based on aspect. Mesic northern aspects ranged from 

315 to 45°, xeric southern aspects ranged from 135 to 

225°, and others were intermediate (Menzel et al. 2006). 

Th ese habitat variables are commonly recorded for small 

mammal and herpetofaunal studies because of their 

infl uence on species occurrence and diversity (e.g., Ford 

et al. 2002, 2006; Francl et al. 2004; Yates et al. 1997).

We examined scatter and residual plots to assess if 

variables met assumptions of analyses (i.e., linearity, 

normality, colinearity). Because all variables deviated 

from normality, we ranked transformed variables and 

performed analysis on the ranked data (Conover and 

Iman 1981a). Untransformed values are presented. 

We standardized amphibian and reptile captures to 

account for diff erences in transect length among gas well 

openings. We compared total reptile, total amphibian, 

red-backed salamander (Plethodon cinereus), Allegheny 

mountain dusky salamander (Desmognathus ochrophaeus), 

small mammal, Peromyscus spp., and meadow vole 

(Microtus pennsylvanicus) captures and total species 

richness and diversity using a two-way analysis of 

covariance with gas well status, aspect, and an interaction 

term as independent variables (Steel and Torrie 1980). 

Because elevation, distance to stream, gas well opening 

area, and road density infl uence small mammal, reptile, 

and amphibian captures (Ford et al. 2002, 2006; Pais 

et al. 1988), and because these variables diff ered among 

sampled gas well sites, we included these as covariates. If 

the analysis of covariance indicated diff erences (P < 0.05), 

we used adjusted Tukey’s Honestly Signifi cant Diff erence 

tests to determine diff erences among independent 

variables. Additionally, we used linear contrasts to 

compare combined storage and producing sites to 

plugged sites to determine diff erences between actively 

maintained and unmaintained well sites. Mean values of 

dependent variables are reported using treatment means 

adjusted for covariates.
5

We compared amphibian, reptile, red-backed 

salamander, and Allegheny mountain dusky salamander 

captures between forest and gas well opening with a twotailed t-test (Conover and Iman 1981a). We compared 

total mammal captures, Peromyscus spp., and meadow 

vole captures among gas well opening, edge, and forest 

with a one-way analysis of variance (Conover and Iman 

1981b). If the analysis of variance indicated diff erences 

(P < 0.05), we used the adjusted Tukey’s Honestly 

Signifi cant Diff erence test to determine individual 

diff erences among independent variables. All statistical 

analyses were conducted using SAS statistical software 

(SAS 2003).

RESULTS

We captured 122 amphibians and 11 reptiles at 40 gas 

well sites, including 66 red-backed salamanders (25 

sites), 35 Allegheny mountain dusky salamanders (15 

sites), 16 slimy salamanders (P. glutinosus; 11 sites), 2 

Wehrle’s salamanders (P. wehrlei; 2 sites), 1 northern 

two-lined salamander (Eurycea bislineata), 1 American 

toad (Bufo americana), 1 eastern red newt (Notophthalmus 

viridescens), 1 eastern garter snake (Th amnophis sirtalis), 

7 ringneck snakes (Diadophus punctatus; 3 sites), 1 redbellied snake (Storeria occipitomaculata), 1 black rat snake 

(Elaphe obsoleta), and 1 eastern fence lizard (Sceloporus 

undulatus). Amphibian, reptile, red-backed salamander, 

and Allegheny Mountain dusky salamander captures 

did not diff er among gas well status types (Table 1). 

Amphibian (F2,27 = 0.74; P = 0.489), reptile (F2,27 = 

0.45; P = 0.643), red-backed salamander (F2,27 = 0.59; 

P = 0.563), and Allegheny Mountain dusky salamander 

(F2,27 = 1.63; P = 0.215) captures did not diff er among 

mesic, xeric, and intermediate aspects. Additionally, we 

did not detect an interaction between gas well status and 

aspect for captures of amphibian (F4,27 = 1.20; P = 0.333), 

reptile (F4,27 = 0.72; P = 0.588), red-backed salamander 

(F4,27 = 0.69; P = 0.603), and Allegheny Mountain 

dusky salamander (F4,27 = 1.27; P = 0.308). Amphibian 

captures increased with increasing road densities (F1,27 

= 4.32; P = 0.047). Amphibian, red-backed salamander, 

and Allegheny Mountain dusky salamander captures 

were greater along forested transects than along gas well 

opening transects (Table 2).

Table 1.—Adjusted means (± SE) for amphibian, reptile, red-backed salamander (Plethodon cinereus), Allegheny 

mountain dusky salamander (Desmognathus ochrophaeus), small mammal, Peromyscus spp., meadow vole (Microtus 

pennsylvanicus) captures and total species diversity (H’) and richness on plugged (inactive; n = 11), storage (n = 16), 

and producing (n = 13) gas well sites and adjacent forest sites within the Monongahela National Forest, West Virginia, 

June–August 2006

Status

 Plugged Storage Producing F2,27

P

Herpetofauna 

 Total amphibians 14.48 ± 3.71 23.89 ± 3.12 21.88 ± 4.22 1.59 0.223

 Total reptiles 18.96 ± 2.91 22.29 ± 2.45 16.92 ± 3.31 0.91 0.414

 Red-backed salamanders 14.32 ± 4.26 25.17 ± 3.59 20.35 ± 4.85 1.54 0.233

 Allegheny Mountain dusky salamander 16.57 ± 3.36 25.99 ± 2.83 22.99 ± 3.82 1.87 0.173

Small Mammals

 Total small mammals 18.97 ± 3.81 20.13 ± 3.21 19.70 ± 4.34 0.02 0.978

 Peromyscus spp. 18.51 ± 3.74 20.36 ± 4.26 20.36 ± 4.26 0.08 0.920

 Meadow vole 18.35 ± 2.71 21.29 ± 2.28 22.63 ± 3.08 0.55 0.585

Shannon-Weiner Diversity Index (H’) 13.50 ± 3.61 24.45 ± 3.04 23.51 ± 4.11 2.45 0.105

Species richness 11.64 ± 3.28A 24.32 ± 2.76B 24.56 ± 3.74AB 4.34 0.023

Means within rows with different letters differ P < 0.05
6

Overall trap success for small mammals was low as we 

captured only 50 individuals over 7,200 trap nights at 

40 gas well sites, including 25 meadow voles (Microtus 

pennsylvanicus; 8 sites), 19 Peromyscus spp. (15 sites), 

2 northern short-tailed shrews (Blarina brevicauda; 2 

sites), 2 southern red-backed voles (Myodes gapperi; 2 

sites), 1 woodland vole (Microtus pinetorum), and 1 

eastern chipmunk (Tamias striatus). Total captures of 

small mammals, Peromyscus spp., and meadow voles did 

not diff er among gas well site status (Table 1). Total 

mammal (F2,27 = 1.87; P = 0.174), Peromyscus spp. (F2,27 

= 1.35; P = 0.276), and meadow vole (F2,27 = 1.70; P = 

0.202) captures did not diff er among mesic, xeric, and 

intermediate aspects. Additionally, we did not detect 

an interaction between gas well status and aspect for 

total number of captures of small mammal (F4,27 = 0.33; 

P = 0.854), Peromyscus spp. (F4,27 = 0.69; P = 0.605), 

or meadow voles (F4,27 = 0.68; P = 0.611). Meadow 

vole captures were negatively related to stream distance 

(F1,27 = 4.37; P = 0.046). Peromyscus spp. captures were 

greater in forested habitat than on gas well sites (Table 

3). Meadow vole captures were greater on gas well sites 

than on forest or edge sites (Table 3).

Combined small mammal and herpetofaunal species 

richness was greater at storage sites than at plugged sites 

(Table 1). Species diversity (F2,27 = 1.10; P = 0.347) and 

richness (F2,27 = 0.69; P = 0.512) did not diff er among 

xeric, mesic, or intermediate aspects. Total species 

diversity was negatively related to stream distance (F1,27 

= 5.33; P = 0.029). Additionally, we did not detect an 

interaction for species diversity (F4,27 = 0.25; P = 0.906) 

and richness (F4,27 = 0.51; P = 0.728). Linear contrasts 

revealed that species diversity (F1,27 = 4.79; P = 0.038) 

and species richness (F1,27 = 8.65; P = 0.007) were greater 

in combined storage and producing sites than in plugged 

sites.

DISCUSSION

Our results support the contention that central 

Appalachian forested habitats harbor greater population 

densities of woodland salamanders than those lacking 

canopy cover (Ash 1997, Ford et al. 2002, Harpole 

and Haas 1999, Petranka et al. 1993). In our study, 98 

percent of salamander captures occurred along forest 

transects, although overall captures were low. During 

daylight hours, microclimatic conditions within forest 

openings are characterized by increased fl uctuations 

in solar radiation, ground temperature, and humidity 

relative to forested habitats (Chen et al. 1999, Geiger 

et al. 1995). Because woodland salamanders require 

moist microclimatic conditions to facilitate effi cient 

 Opening Edge Forest F P

Total small mammals 0.65 ± 0.36 0.30 ± 0.11 0.40 ± 0.11 0.32 0.728

Peromyscus spp. 0.03 ± 0.03A 0.15 ± 0.07AB 0.35 ± 0.10B 5.65 0.005

Meadow voles 0.60 ± 0.36A 0.05 ± 0.05B 0.00 ± 0.00B 6.18 0.003

Table 3.—Mean (± SE) captures of small mammals, Peromyscus spp., and meadow voles (Microtus pennsylvanicus) 

within gas well opening, edge, and forested habitat at gas well sites (n = 40) within the Monongahela National Forest, 

West Virginia, June-August 2006

Means within rows with different letters differ P < 0.05

Table 2.—Mean (± SE) captures of amphibians, reptiles, red-backed salamanders (Plethodon cinereus), and 

Allegheny Mountain dusky salamanders (Desmognathus ochrophaeus) along transects in gas well opening and 

forested habitat at gas well sites (n = 40) within the Monongahela National Forest, West Virginia, June-August 2006

 Forest Gas well t P

Amphibians 0.030 ± 0.004 0.002 ± 0.001 8.26 <0.001

Reptiles 0.001 ± 0.000 0.007 ± 0.003 -0.88 0.384

Red-backed salamanders 0.017 ± 0.003 0.000 ± 0.000 7.76 <0.001

Allegheny Mountain dusky salamanders 0.009 ± 0.002 0.000 ± 0.000 4.80 <0.001
7

cutaneous respiration (Spotila 1972), moisture and 

temperature fl uctuations may surpass their tolerance 

thresholds within gas well openings. However, suffi cient 

rock and CWD cover availability may mitigate adverse 

microclimatic fl uctuations by providing moist refugia 

(Marsh et al. 2004, Riedel et al. 2008, Young and 

Yahner 2003). Riedel et al. (2008) observed frequent 

red-backed salamander occurrence within unmowed 

hay meadows adjacent to woodlots in southern West 

Virginia, attributing their occurrence within this habitat 

to cover object availability retained within meadows. 

Although we did not quantify cover object abundance 

within gas well openings and forest habitats, we observed 

signifi cantly fewer large pieces of CWD or rock within 

openings. Th erefore, if suffi cient cover availability was 

retained within gas well openings during development, 

it may mitigate, to some extent, adverse microclimatic 

conditions.

Reduced woodland salamander abundance along forest 

edges has been attributed to unfavorable microclimatic 

conditions from adjacent forest openings, particularly 

those created from timber harvest (DeGraaf and 

Yamasaki 2002, deMaynadier and Hunter 1998) 

and roads (Marsh and Beckman 2004, Marsh 2007, 

Semlitsch et al. 2007). However, we found greater 

amphibian captures with increasing road densities, 

providing additional support for lack of adverse edge 

eff ects on woodland salamanders observed by Moseley et 

al. (2009), where woodland salamander occurrence was 

greater within 60 m of gas well opening edge. Similarly, 

DeGraaf and Yamasaki (2002) observed increasing redbacked salamander abundance 5 to 20 m from edge into 

mature northern hardwood forests in New Hampshire. 

Moreover, we observed the presence of American toad 

tadpoles within many roadside ditches containing 

standing water adjacent to well openings, suggesting 

these areas may serve as breeding habitat (Barry et al. 

2008, Cromer et al. 2002).

Contrary to our initial hypothesis, reptile capture rates 

did not diff er among forest and gas well openings. 

However, because reptiles are ectothermic, they are 

dependent on habitat attributes, such as forest canopy 

openings, that enable effi cient body temperature 

regulation to facilitate optimal foraging, breeding, 

and predator avoidance behavior (Seigel et al. 1987). 

Ringneck snakes were the most common reptile 

captured (n = 7) on our gas well sites. Although ringneck 

snakes often are associated with mesic forest habitat 

in southeastern Piedmont and Coastal Plain regions 

(Willson and Dorcas 2004), 85 percent of our captures 

occurred within gas well openings. Our sampling 

techniques may have been insuffi cient for sampling 

reptile species, particularly lizards and large snakes (Ryan 

et al. 2002), and small sample size precludes strong 

inference regarding gas well opening use by reptiles. 

Nonetheless, because the MNF is 97 percent forested, 

experiences cooler, moister conditions relative to more 

xeric Appalachian regions, and has extended periods 

below optimal temperature thresholds for reptile species 

(Seigel et al. 1987, U.S. Forest Service 2006), gas well 

openings may serve as important basking and egg-laying 

habitat (Kjoss and Litvaitus 2001). Pais et al. (1988) 

reported greater herpetofauna richness in 0.2- to 1.0-ha 

wildlife openings relative to mature hardwood forest 

in Kentucky. As with woodland salamanders, large 

cover items within openings may have been limiting 

for smaller snakes. Because our sampling occurred 

during summer, when temperatures were likely above a 

threshold maximum for basking, reptiles may not use gas 

well openings as much as during fall and spring when 

temperatures are cooler. Further seasonal investigations 

examining reptile use of gas well openings and other 

early successional habitats in the central Appalachian 

mountain region are needed.

In the central Appalachian Mountains, meadow voles 

are strongly associated with old-fi eld and meadow 

habitats (Ford et al. 2007, Francl et al. 2004). Indeed, all 

meadow vole captures in our study occurred within gas 

well openings. Moreover, 56 percent of all meadow vole 

captures occurred at one site where a thick debris layer 

was present, which may also account for the observed 

negative relationship between increasing meadow vole 

captures and decreasing stream distance; this particular 

well site was located only 573 m from a stream. Meadow 

voles can be abundant where a thick layer of grass 

provides greater cover (M’Closkey and Fieldwick 1975). 

Overall, meadow vole captures in our study were low 
8

and therefore conclusions about gas well opening habitat 

quality must be made with caution. Also, semi-annual 

mowing, in addition to soil compaction within well 

openings (Adams 2007, Moseley et al. 2009), may be 

suppressing establishment of suitable vegetative cover, 

and thus reducing meadow vole abundance. Inadequate 

vegetative cover may increase predation risk from mesomammalian predators that frequently hunt in forest 

openings (Parker et al. 1992).

Many of our sampled gas well openings may not be large 

enough to support viable meadow vole populations. 

Despite their preponderance, we successfully trapped 

meadow voles at only 8 of 40 sites sampled. In West 

Virginia, meadow voles were generally absent from early 

successional wetland habitats less than 1.3 ha (Francl 

et al. 2004). Furthermore, Francl et al. (2004) found 

meadow vole captures were greater at wetlands with 

increased road density. Similarly, small mammal species 

associated with early successional habitats occurred at 

higher densities in larger clearcut openings (> 25 ha) than 

smaller openings (< 6 ha) within pine (Pinus spp.) stands 

in the Upper Coastal Plain of South Carolina (Yates et al. 

1997). Because lower density small mammal populations 

within small habitat patches are more susceptible to 

local extinction, migration corridors that facilitate 

recolonization and augmentation of existing populations 

within patches are necessary for long-term persistence 

(Menzel et al. 2005, Yates et al. 1997). Indeed, Menzel 

et al. (2005) attributed skid roads associated with timber 

harvest to increased small mammal richness and diversity 

within bottomland hardwood forest gaps in the South 

Carolina Coastal Plain. We did not sample adjacent 

roads and gas pipeline rights-of-way, which likely serve as 

important migration routes for meadow voles and other 

small mammals (Cummings and Vessey 1994, Francl 

et al. 2004, Getz et al. 1978) within the predominately 

forested MNF landscape.

In the eastern U.S., Peromyscus spp. occur in a variety 

of habitats with particular affi nity for areas with diverse 

vegetative structure (Dueser and Shugart 1978, Myton 

1974, Osbourne et al. 2005). We captured Peromyscus

spp. more often in edge and forest habitats than in gas 

well openings. Similarly, deer mouse captures increased 

with increasing distance from wildlife openings within 

northern hardwood forests in the southern Appalachians 

of North Carolina (Menzel et al. 1999). Although not 

quantifi ed, edge habitats at most of our sites had welldeveloped vegetative structure or “side-canopy” (Didham 

and Lawton 1999, Matlack 1994) that may account for 

greater numbers of Peromyscus spp. captures within edge 

and forest interior habitats. However, our low sample 

size precluded examination of demographic parameters 

between habitat types, thereby limiting inferences about 

habitat quality associated with gas well sites (Van Horne 

1983).

Our results suggest maintained gas well sites support 

greater species richness and diversity than nonmaintained plugged well sites. Although not maintained, 

plugged wells had greater soil compaction than in storage 

well openings (Adams 2007), which may have reduced 

long-term availability of underground burrow habitat for 

small mammals and herpetofauna. Additionally, because 

plugged sites are no longer mowed, these sites had greater 

tree cover than storage or producing wells (Adams 2007). 

Th erefore, plugged sites may have provided less suitable 

basking sites to reptile species and inadequate grass/forb 

structure to meadow voles as succession has transitioned 

these sites beyond old-fi eld habitats.

CONCLUSION

Maintained forest openings and other early successional 

habitats are among the most rapidly declining habitats 

throughout the Appalachian Mountains (Litvaitis 2001). 

Recruitment and maintenance of early successional 

habitat have declined as farm abandonment progresses 

to forest cover, and as fi re suppression increases, and 

even-aged timber harvesting is increasingly limited 

(Trani et al. 2001). Although gas well development 

represents a more permanent form of disturbance 

relative to natural perturbations, our results suggest 

that gas well openings can contribute to increased 

species richness and diversity within predominately 

forested central Appalachian landscapes. Additionally, 

natural gas development generally requires less land 

disturbance than development for other fossil fuels, 

such as petroleum and coal (U.S. Department of Energy 

2003), thereby minimizing its impact on forest interior 
9

associated species. Th e utility of gas well openings as 

wildlife habitat may be further improved by adhering 

to West Virginia Best Management Practice guidelines, 

particularly with regard to alleviating soil compaction 

(West Virginia Department of Natural Resources 

2006) to facilitate burrowing and development of dense 

vegetation. Furthermore, management of gas well sites 

as wildlife openings will benefi t popular game species 

such as the eastern wild turkey (Meleagris gallopavo), 

American black bear (Ursus americanus), and whitetailed deer (Odocoileus virginianus; Kammermeyer 

and Moser 1990, Parker et al. 1992, Wentworth et al. 

1990). Wildlife opening maintenance for game species 

also benefi ts numerous songbird species by increasing 

habitat heterogeneity and food resources (Parker et al. 

1992). Investigations regarding vegetation manipulation 

through altered mowing regimes and increasing cover 

item (e.g., CWD and rocks) availability within gas well 

openings are needed to facilitate a better understanding 

of gas well openings as early successional habitat in the 

central Appalachian Mountains. Also, more research is 

needed on other wildlife species with varying habitat 

requirements before attributing overall wildlife benefi ts 

to natural gas development. In particular, the potential 

for negative impacts to aquatic biota from surface 

and groundwater contamination and to cave-dwelling 

organisms where karst geology is encountered (Goodbar 

1997) requires further examination. Ongoing natural gas 

development on the Monongahela National Forest in 

karst geology and on the Allegheny National Forest off ers 

opportunities for such investigations. (U.S. Forest Service 

2007).

ACKNOWLEDGMENTS

We thank L. Tracy for logistical support and M. Strager 

for GIS support. Funding for this study was provided 

by the USDA Forest Service Monongahela National 

Forest and the Northern Research Station and the West 

Virginia University Division of Forestry and Natural 

Resources through participating agreement No. 06–PA–

11092100–041.

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Moseley, Kurtis R.; Ford, W. Mark; Edwards, John W.; Adams, Mary B. 2010. 

Reptile, amphibian, and small mammal species associated with natural gas 

development in the Monongahela National Forest, West Virginia. Res. Pap. 

NRS-10. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, 

Northern Research Station. 14 p.

Burgeoning energy demand in the United States has led to increased natural gas 

exploration in the Appalachian Basin. Despite increasing natural gas development in 

the region, data about its impacts to wildlife are lacking. Our objective was to assess 

past and ongoing natural gas development impacts on reptiles, amphibians, and small 

mammals in the Monongahela National Forest in West Virginia. We sampled 40 gas 

well sites and compared amphibian, reptile, and small mammal captures among active 

producing, plugged (inactive), and storage well types. Total species richness and 

diversity were greater at storage gas well sites than at plugged wells. Although natural 

gas development adversely impacts moisture-sensitive woodland salamanders, our 

results suggest that maintained gas well openings may benefi t other herpetofauna 

and small mammal species that use early successional habitat within predominately 

forested central Appalachian landscapes.

KEY WORDS: Allegheny Mountains, Microtus, natural gas, Peromyscus, Plethodon 

cinereus
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