nena masthead
SENA Home Staff & Editors For Readers For Authors

Bats of the Cumberland Plateau and Ridge and Valley Provinces, Virginia
John Timpone, Karen E. Francl, Dale Sparks, Virgil Brack, Jr., and Joel Beverly

Southeastern Naturalist, Volume 10, Issue 3 (2011): 515–528

Full-text pdf (Accessible only to subscribers.To subscribe click here.)


Access Journal Content

Open access browsing of table of contents and abstract pages. Full text pdfs available for download for subscribers.

Issue-in-Progress: Vol. 22 (2) ... early view

Current Issue: Vol. 21 (4)
SENA 21(3)

All Regular Issues


Special Issues






JSTOR logoClarivate logoWeb of science logoBioOne logo EbscoHOST logoProQuest logo

2011 SOUTHEASTERN NATURALIST 10(3):515–528 Bats of the Cumberland Plateau and Ridge and Valley Provinces, Virginia John Timpone1,*, Karen E. Francl2, Dale Sparks1, Virgil Brack, Jr.1, and Joel Beverly3 Abstract - The distribution and abundance of bats in western Virginia are poorly documented, especially in summer. Herein, we report results for captures of bats during 8 summers (between 2000 and 2009) of mist-netting surveys in the Cumberland Plateau and the Ridge and Valley provinces of Virginia. We captured 1575 bats representing 11 species and report multiple new county records for 10 of the 11 counties surveyed. Average capture rate was 1.9 bats per net-night, and Simpson’s diversity index was 3.9 for the 8-year period. We also documented earliest and latest detection dates for reproducing females and presence of juveniles in 7 bat species. Our data contribute to an understanding of the relative abundance, distribution, and reproductive phenology of bats in these provinces, and may aid in the development of sound conservation strategies for these species in Virginia and surrounding states. Baseline data like these are increasingly important in light of White-Nose Syndrome’s effects on bat populations in the region. Introduction Although bats play important ecological (e.g., Duchamp et al. 2010, Fenton 1997) and economic (Cleveland et al. 2006) roles, conservation is hindered by gaps in information, including a lack of fundamental data on distribution and abundance of species across their respective ranges. Studies of species’ distributions provide critical baseline data that can be compared with future datasets to help determine how and why distributions change over space and time (Menzel et al. 2003, Sparks and Choate 2000). Regional data are particularly important, as ecological and anthropogenic changes, such as White-Nose Syndrome (WNS; Blehart et al. 2009, Boyles and Willis 2009) and wind turbines (Arnett 2005, Kunz et al. 2007), threaten to alter regional bat communities. Seventeen species of bats are known from Virginia (Linzey 1998), 14 of which have been previously documented in the Cumberland Plateau or Ridge and Valley provinces (Fig. 1) in western Virginia (Hobson 1998). Four of these species are listed as federally or state endangered, including Myotis sodalis Miller and Allen (Indiana Bat), Myotis grisescens Howell (Gray Bat), Corynorhinus townsendii virginianus Handley (Virginia Big-eared Bat), and Corynorhinus rafinesquii macrotis Lesson (Southeastern Big-eared Bat), and 2 are considered rare (Myotis leibii Audobon and Bachman [Eastern Small-footed Bat] and Myotis austroriparius Rhoads [Southeastern Bat]; Roble 2006). Despite this diversity, published data on abundance and distribution of bats in western Virginia are limited to winter studies or field guides and similar texts (e.g., Bailey 1946, Linzey 1998, 1Environmental Solutions and Innovations, Inc. Cincinnati, OH 45233. 2Biology Department, Radford University, Radford, VA 24142. 3Apogee Environmental, Whitesburg, KY 41858. *Corresponding author - 516 Southeastern Naturalist Vol. 10, No. 3 Webster et al. 1985, Whitaker and Hamilton 1998). The most complete dataset consists of winter hibernacula surveys in which hibernating bats were monitored in 170 caves west of the Blue Ridge Mountains over a 10-year period (Dalton 1987), an effort that has been sustained and intensified by the Virginia Department of Game and Inland Fisheries (VDGIF; Terwilliger 1991). However, quantitative summer data remain limited. Herein we provide results for 8 summers of mist-netting surveys in the Cumberland Plateau and the Ridge and Valley provinces of Virginia. These data contribute to an understanding of relative abundance, distribution, and reproductive phenology of bats, and may aid development of sound conservation strategies for these species in Virginia and the region. Furthermore, in light of the decimating impact of White-Nose Syndrome on bat populations documented in the northeastern United States (Blehart et al. 2009), such baseline data provide a decade-long snapshot of bat communities by which we can measure species-specific temporal changes in relative abundance and reproductive status in these provinces. Methods Study area We conducted surveys in 11 counties (Bland, Buchanan, Dickenson, Pulaski, Rockingham, Russell, Shenandoah, Smyth, Tazewell, Wise, and Wythe) wholly or partially within the Cumberland Plateau and Ridge and Valley physiographic provinces of western Virginia (Fig. 1). The Cumberland Plateau is characterized by steep slopes interspersed with gently sloping plateau remnants and numerous streams. The Ridge and Valley province to the East contrasts the Cumberland Plateau by having longitudinal ridges and steeper depressions in strata (Fenneman 1938). Dominant vegetation of this region prior to settlement was characterized by mixtures of Quercus spp. (oaks) and Castanea dentata (Marsh.) Borkh. (American Chestnut), with pockets of mixed mesophytic forest in coves, ravines, and other fertile sites (Braun 1950). Following elimination of American Chestnut as an overstory tree by about 1940, this region is now mostly described as mixed oak forest (Stephenson 1993). Figure 1. Counties surveyed for bats between 2000–2001 and 2004–2009 in the Cumberland Plateau (counties 1–3) and the Ridge and Valley (counties 4–11) physiographic provinces of Virginia. 2011 J. Timpone, K.E. Francl, D. Sparks, V. Brack, Jr., and J. Beverly 517 Survey techniques Mist-net surveys were conducted between 15 May to 15 August 2000–2001 and 2004–2009 at 201 sites for a total of 402 nights (804 complete net-nights). A complete night of netting lasted for at least 5 hours, typically beginning at dusk or shortly thereafter. Netting was suspended during adverse weather (temperatures <10 °C, strong winds, or precipitation) in order to comply with guidelines for sampling for the Indiana Bat (USFWS 1999, 2007). Data from partial net-nights were not included in the analysis. We captured Bats using double- (5.2 m high) or triple- (7.8 m high) stacked mist-nets. Nets typically were set across forested corridors and first- or second-order streams used as travel corridors by bats. Width of nets varied from 6.0–18.0 m, depending on width of the corridor. We identified bats to species, and recorded sex, reproductive condition, age (juvenile or adult, based on degree of epiphyseal ossification [Anthony 1988]), length of right forearm, and mass. Pregnancy was determined by palpitation of the abdomen, and status of lactation was determined by condition of teats and the ability to exude milk (Racey 1988). Documentation and analyses We determined if occurrences represented new county records using the Biota of Virginia database, which is supported by reports made by individuals with scientific collecting permits (VGDIF 2011). Chi-square analysis was used to compare overall capture rates of adult males and adult females. Simpson’s reciprocal index was used to quantify diversity (D), where D = l / ΣPi 2, and Pi is the proportion of bats belonging to species i (MacArthur 1972). This index has been used in similar studies of bats (e.g., Brack 2009) and was used here to allow comparisons. Results We captured 1575 bats of 11 species. Five species comprised 93% of all captures and were documented in at least 9 counties, including: Eptesicus fuscus Beauvois (Big Brown Bat; 28% of all individuals captured), Myotis septentrionalis Trouessart (Northern Bat; 21%), Lasiurus borealis Müller (Eastern Red Bat; 17%), Myotis lucifugus LeConte (Little Brown Bat; 16%), and Perimyotis subflavus Menu (Tri-colored Bat; 11%). The remaining six species, Eastern Small-footed Bat (5%), Lasionycteris noctivagans LeConte (Silver-haired Bat; 0.9%), Lasiurus cinereus Beauvois (Hoary Bat; 0.5%), Gray Bat (0.4%), Virginia Big-eared Bat (0.1%), and Indiana Bat (0.1%), were captured less frequently and documented in only 1–5 counties. Mean rate of capture was 1.9 bats per net-night (range = 1.2 to 3.4). Simpon’s diversity index across all years was 3.9 and ranged from 3.1 to 4.8. Overall, we captured significantly more adult males (n = 598) than adult females (n = 501; χ2 = 8.56, P = 0.001). Within species, we captured significantly more adult male Red (χ2 = 59.58, P < 0.001), Silver-haired (χ2 = 13.00, P = 0.004), and Tri-colored Bats (χ2 = 52.57, P < 0.001) than adult females. Conversely, we captured significantly more adult female Northern (χ2 = 6.93, P = 0.008) and Big 518 Southeastern Naturalist Vol. 10, No. 3 Brown Bats (χ2 = 4.14, P = 0.042) than adult males. Evidence of reproduction was observed for 7 of the 11 species (Table 1). Throughout the study, we captured 90 pregnant females representing 7 species. Pregnant females were captured as early as 15 May (first day of sampling) and as late as 24 June. We also captured 131 lactating females representing 6 species. Earliest and latest capture dates of lactating females were 29 May and 22 July, respectively (Table 1). For all species, we documented 476 juveniles (30% of all captures); the earliest date of capture for a juvenile bat was 16 June, and the latest was 15 August (last day of sampling). Discussion Although we documented new records of species in 10 of 11 counties surveyed, it is unlikely that these represent range extensions. Rather, our captures likely fill gaps in data. The rate of captures in the present study was higher than those reported from West Virginia by Brack et al. (2005; 1.4 bats per net-night, diversity = 4.0) and Menzel et. al. (2002; 0.76 bats per net-night, diversity not reported). Compared to previous studies in other eastern states, our results yielded lower capture success but higher diversity. In northern Pennsylvania, Brack (2009) reported a capture rate of 2.9 bats per net-night and diversity of 3.2. The capture rate and diversity from a similar study in Ohio was 2.4 bats per net-night and 2.9, respectively (Brack and Duffey 2006). Below, we report species-specific trends in distribution and reproductive status. Species are presented according to federal or state protection status, as classified in the Biota of Virginia database. Non-game, protected species Big Brown Bat. The Big Brown Bat is considered a habitat generalist (Agosta 2002) and is one of the most widespread and common species in North America. Big Brown Bats were the most frequently encountered species (28% of all individuals), with an overall capture rate of 0.55 bats per net-night (range = 0.28 to 1.01). Big Brown Bats comprised 16%, 18%, and 35% of total captures in past surveys in the Appalachians of Virginia (Hobson 1998), West Virginia (Menzel et al. 2002), and North Carolina (O’Keefe 2009), respectively. Sampling bias may partially explain the higher capture rate of adult females Table 1. Dates of capture for reproductive female (n = 469) and juvenile bats (n = 187) in 11 counties in the Cumberland Plateau and Ridge and Valley physiographic provinces of Virginia, 2000–2001 and 2004–2009. Dates are the earliest and latest dates of capture during the standardized surveying season of 15 May–15 August. “N.D.” indicates no bats were caught. Pregnant Lactating Post-lactating Juvenile Species Earliest Latest Earliest Latest Earliest Latest Earliest Latest Big Brown Bat 15 May 13 June 2 June 17 July 20 June 15 Aug 16 June 15 Aug Red Bat 15 May 8 June 29 May 11 July 14 July 14 Aug 24 June 15 Aug Hoary Bat 25 June 21 July N.D. N.D. 25 June 21 July 25 July 15 Aug E. Small-footed Bat 18 May 20 June 17 June 27 June 8 Aug 13 Aug 29 July 9 Aug Little Brown Bat 6 June 22 June 21 June 18 July 14 July 15 Aug 20 June 14 Aug Northern Bat 17 May 19 June 17 June 22 July 23 June 10 Aug 10 July 14 Aug Tri-colored Bat 22 May 24 June N.D. N.D. 15 Aug 15 Aug 7 Aug 12 Aug 2011 J. Timpone, K.E. Francl, D. Sparks, V. Brack, Jr., and J. Beverly 519 than adult males. Net sites over riparian areas, often thought to be foraging hot spots for adult females (Barclay 1989b, Grindal et al.1999), were sampled more frequently than other habitats. In addition, lactating females make more drinking passes than non-reproductive females (Adams and Hayes 2008). Evidence of reproduction was documented in all counties surveyed, indicating the presence of maternity colonies. While maternity colonies may contain up to several hundred adult females, adult males usually are not typically associated with maternity colonies and appear to be more dispersed (Whitaker and Hamilton 1998). Parturition occurred as early as 2 June and volant young first appeared as early as 16 June. In a building in southern Michigan, lactating Big Brown Bats were reported as early as 3 June and volant young by 23 June (Kurta 2010). Northern Bat. The Northern Bat is a forest-obligate species that may be most abundant in the forest-dominated landscape of the Appalachian Mountains, and our capture rates are consistent with other studies from the region (Brack 2009, Brack et al. 2005, Castleberry et al. 2007, Menzel et al. 2002). Northern Bats were the second most abundant species captured (21% of all captures; 50% of all Myotis spp.) during our surveys, with an average capture rate of 0.41 bats per net-night (0.14 to 0.50). We obtained 5 new county records for this species (Buchanan, Pulaski, Shenandoah, Smyth, and Wise). Although we captured more adult females than adult males, we attribute these results to sampling bias (see above). While females usually form maternity colonies in trees, harboring between 30–50 bats (Carter and Feldhamer 2005, Foster and Kurta 1999, Timpone et al. 2010), males are typically more dispersed and roost either singly or in small groups in trees, caves, mines, or manmade structures (Agosta et al. 2005, Broders and Forbes 2004, Perry and Thill 2007). Extending our sampling period, which began when maternity colonies had already been established and ended when maternity colonies were beginning to disband, may have yielded a more even adult sex ratio. Pregnant Northern Bats were documented between 17 May and 19 June, and lactating bats were documented from 17 June to 22 July. The earliest date of capture of a juvenile Northern bat was 20 July. Similarly, Handley (1991) found this species can comprise 35% or more of Myotis species in late summer. Red Bat. Red Bats are common summer residents throughout much of the eastern United States, including the Appalachian Mountains (Hutchinson and Lacki 1999, Menzel et al. 2002, O’Keefe 2009). Red Bats are seasonal migrants (Whitaker and Hamilton 1998), and although there are reports of individuals overwintering in Ohio and West Virginia (Linzey 1998), it is unknown whether they occur year-round in Virginia. Average capture rate for Red Bats was 0.34 bats per net-night (0.05 to 0.94), and 5 new county records were obtained (Buchanan, Dickenson, Pulaski, Russell, and Tazewell). Male Red Bats comprised over 80% of adult captures. The paucity of reproductive females captured during our surveys is consistent with other studies conducted in the Appalachian Mountains of North Carolina (O’Keefe 2009) and West Virginia (Brack 2009, Brack et al. 2005, Menzel et al. 2002). Males accounted for 93% and 95% of all Red Bat captures in Virginia (Hobson 1998) and West Virginia (Castleberry et al. 2007), respectively. Ford et al. (2002) suggest that female Red Bats are more abundant 520 Southeastern Naturalist Vol. 10, No. 3 in the Coastal Plain than the Appalachian Highlands, due to warmer summer temperatures at lower elevations. In support of this idea, the majority of Red Bats captured by Johnson and Gates (2008) on Maryland’s Coastal Plain were reproductive females and juveniles. Pregnant Red Bats were taken 15 May through 8 June, and lactating Red Bats were captured from 29 May though 11 July. The earliest capture date for a volant juvenile was 24 June. In Michigan’s Lower Peninsula, pregnant Red Bats were captured from 29 May to 28 June, lactating bats were reported from 15 June to 4 August, and earliest date of capture for juveniles was 2 July (Kurta 2010). Little Brown Bat. The Little Brown Bat is one of the most widespread species in North America, but varies in local abundance. It was the fourth most commonly encountered species during our surveys (16% of all individuals), with an average capture rate of 0.30 bats per net-night (0.06 to 0.54). We caught this species in six counties, two of which (Buchanan and Shenandoah) were new county records. The Little Brown Bat was the most frequently captured species reported by Castleberry et al. (2007) in West Virginia and Hobson (1998) in Virginia. Pregnant females were captured between 6 June and 22 June, while lactating females were reported between 21 June and 18 July. Our earliest capture of a juvenile Little Brown Bat was 20 June. Kurta (2010) reported that Little Brown Bats from a maternity colony in a building in southern Michigan gave birth as early as 6 June and young were flying as early as 23 June. Tri-colored Bat. The Tri-colored Bat is widespread throughout much of the eastern United States (Fujita and Kunz 1984) and is considered one of the most common bats in Virginia (VDGIF 1988). We captured this bat in 10 of the 11 survey counties, including two new county records (Wythe, Dickenson) at a rate of 0.21 bats per net-night (0.04 to 0.70). The male-biased sex ratio we observed seems to be typical of the Appalachian region. In a study of 11 mid-Atlantic National Parks in four physiographic provinces, 94% (n = 16) of adult female Tri-colored Bats were captured in the Coastal Plain and Piedmont, while only 6% (n = 1) were captured in the Blue Ridge and Ridge and Valley physiographic provinces (Johnson et al. 2008). In western Virginia, 88% (n = 44) of all Tri-colored Bats captured were males (Hobson 1998). Similarly, males comprised 94% all captures of Tri-colored Bats (n = 17) in West Virginia (Castleberry et al. 2007). Pregnant females were captured between 22 May and 24 June, and earliest capture of a juvenile was 7 August. No data were available for lactating females. While comparative data on reproductive timing for this species was not available for the region of our study, Veilleux et al. (2004) reported pregnant females between 6 May and 23 June, lactating females between 29 June and 6 July, and volant young as early as 16 July in Indiana. Silver-haired Bat. VDGIF’s records of Silver-haired Bats are limited to a few scattered counties, although this species is listed as “likely” throughout the Commonwealth (Linzey 1998). We captured this species at an average rate of 0.01 bats per net-night (0 to 0.07). Captures in Buchanan and Tazewell counties were new county records. Low capture success but high likelihood of county presence may be related to their morphology and foraging habits. The Silver-haired Bat 2011 J. Timpone, K.E. Francl, D. Sparks, V. Brack, Jr., and J. Beverly 521 has lower wing-aspect ratio, higher wing-loading, and lower call frequencies than most species of Myotis (Aldridge and Rautenbach 1987), making it more suited for foraging in open habitats than in the heavily forested stream and road corridors we sampled. In the central Appalachians of West Virginia, Owen et al. (2004) documented higher rates of activity of Silver-haired Bats in relatively uncluttered habitats such as clear-cuts. All of the Silver-haired Bats we captured were adult males, which is consistent with Hobson’s (1998) capture of 3 adult males and Cryan’s (2003) study of gender segregation during the summer months. Kurta (2010) reported that males comprised 88% of adult Silver-haired Bats captured in Michigan over 32 years, and Kunz (1982) reported similar seasonal gender segregation throughout this species’ eastern range. Apparently, females migrate to the northern United States and Canada to raise young, while males remain nearer to their winter range (Cryan 2003). Hoary Bat. The Hoary Bat is listed as “likely” throughout the Commonwealth (Linzey 1988). However, each of the 6 counties where Hoary Bats were captured is a new county record. Hoary Bats comprised a small fraction of total captures (0.009 bats per net-night). This result was similar to other studies conducted in the Appalachians, where the species accounted for 0.005 percent of total captures in West Virginia (Castleberry et al. 2007) and 0.004 percent of total captures in Virginia (Hobson 1998). Although Hoary Bats were infrequently captured, they are probably more widespread than our data indicate. Like Silver-haired Bats, Hoary Bats possess both morphological and acoustical traits suited for foraging in open habitats (Aldridge and Rautenbach 1987, Barclay 1989a), and they frequently fly at heights in excess of 50 m (Barbour and Davis 1969). Using nets up to 20 m high, Brack (1985) found that 50% of Hoary Bat captures were at heights >8.3 m. Furthermore, in the central Appalachians of West Virginia, Owen et al. (2004) detected a greater number of echolocation calls of Hoary Bats in open-canopied habitats (e.g., clear-cuts) than in closed-canopied habitats. Because the majority of our sites were on streams or road corridors with closed canopies and nets were 5.2 to 7.8 m high, this species likely was not adequately sampled by our surveys. Federally endangered species Virginia Big-eared Bat. The capture of a single Virginia Big-eared Bat in Tazewell County suggests minimal species activity in summer. Dalton (1987) found this species hibernating in caves in 5 counties (Tazewell, Bland, Bath, Highland, Rockingham). R. Reynolds (VDGIF,Verona, VA, pers. comm.), W. Orndorff (Virginia Department of Conservation and Recreation, Christiansburg, VA, pers. comm.), and K. Francl (pers. observ.) confirmed the continued presence of this species outside caves during fall swarming in 2009 in Bland and Tazewell counties. The only known maternity colony for this species in Virginia is in Burke’s Garden, Tazewell County (V. Brack, Jr., pers. observ.). Gray Bat. In Virginia, previous records of the Gray Bat during summer indicate this species is restricted to just 4 caves in Lee and Scott counties (not surveyed 522 Southeastern Naturalist Vol. 10, No. 3 in this project). These caves were thought to house only males (Handley 1991, Holsinger 1964). Our discovery of 4 adult males in Russell County support the idea that caves in Virginia mainly support bachelor colonies, and extends knowledge of distribution northeast to a third county. More recent surveys (August 2010 ) by R. Reynolds, W. Orndorff, and K. Francl (pers. observ.) also extend the range of the Gray Bat to 5 bachelor caves in Lee and Scott counties, and a single maternity colony near Bristol, VA. Indiana Bat. Although Indiana Bats are considered more widespread in Virginia than Gray Bats, the species was thought to comprise just 1% of all bats in the Commonwealth (Handley 1991). The winter occurrence of Indiana Bats in 8 caves across 5 of our study counties (Lee, Wise, Tazewell, Bland, Bath), combined with our capture of a single male in Tazewell County, suggests that summer populations may be smaller than expected in the Cumberland Plateau and Ridge and Valley provinces. However, given that male Indiana Bats often remain close to their winter hibernacula during summer (Brack 1983, Whitaker and Brack 2002), our survey may have underestimated their abundance. Federal species of concern Eastern Small-footed Bat. The Eastern Small-footed Bat is listed as both a federal species of concern and a species of concern for the national forests (George Washington-Jefferson) that comprise over 7285 ha concentrated in the western portion of the state and encompassing many of our study sites (USDAFS 2004). We captured an average of 0.09 (0.01 to 0.30) Eastern Small-footed Bats per net-night and documented one new record for Buchanan County. Little is known concerning this species, but recent research indicates that Small-footed Bats may be dependent on rock outcrops for summer roosting habitat (Johnson and Gates 2008) and on surrounding forests for foraging habitat (Johnson et al. 2009). In some southwestern portions of the Commonwealth, Small-footed Bats comprise ca. 15% of Myotis inhabiting caves in late summer (Handley 1991, Handley et al. 1979). Rafinesque’s Big-Eared Bat. Rafinesque’s Big-eared Bat was notably absent from our captures. The disjunct subspecies C. rafinesquii macrotis is listed as a state-endangered species, and is restricted to southeastern Virginia in the Coastal Plain physiographic region. However, C. r. rafinesquii have been captured in Pike County, KY (adjacent to Buchanan County, VA; K. Francl, pers. observ.), and Hancock County, TN (adjacent to Lee and Scott counties [not surveyed]; Handley et al. 1979). Hence, this species is considered “likely” to be present in Lee, Scott, and Washington counties (BOVA database), which border 3 of the counties we surveyed (Wise, Russell, Smyth). Hobson (1998) did not capture any Rafinesque’s Big-eared Bats in western Virginia. However, this species is adept at eluding mist-nets, and the use of mist-nets alone is not an effective sampling protocol (Lance and Garrett 1997). We suggest additional efforts in Virginia are needed, including surveys of abandoned buildings, rock shelters, caves, and bridges throughout the study area. 2011 J. Timpone, K.E. Francl, D. Sparks, V. Brack, Jr., and J. Beverly 523 Conclusion Since the collection of our data, WNS has been confirmed in multiple hibernacula in 6 of the 11 counties in our study, and in 10 of the counties within the physiographic provinces we investigated (R. Reynolds, W. Orndorff, and K. Francl, unpubl. data). Furthermore, wind turbines are proposed for Highland County (Ridge and Valley Province), and are under consideration for Wise and Tazewell counties (Dominion Press Release, in litt.). With multiple studies seeking to quantify the impacts of these major changes (e.g., Dzal et al. 2010;W.M. Ford, Virginia Tech, Blacksburg, VA, pers. comm), our data are an increasingly valuable resource to researchers and resource managers. Acknowledgments We thank the employees of Environmental Solutions and Innovations, Inc. involved in the collection of these data over the years, especially A. Mann, J. Duffey, D. Cox, D. Jeffcott, J. Hootman, and L.M. Gilley. Literature Cited Adams, R.A., and M.A. Hayes. 2008. Water availability and successful lactation by bats as related to climate change in arid regions of western North American. Journal of Animal Ecology 77:1115–1121. Agosta, S.J. 2002. Habitat use, diet, and roost selection by the Big Brown Bat (Eptesicus fuscus) in North America: A case for conserving an abundant species. Mammal Review 32:179–198. Agosta, S.J., D. Morton, B.D. Marsh, and K.M. Kuhn. 2005. Nightly, seasonal, and yearly patterns of bat activity at night roosts in the central Appalachians. Journal of Mammalogy 86:1210–1219. Aldridge, H.D.J.N., and I.L. Rautenbach. 1987. Morphology, echolocation, and resource partitioning in insectivorous bats. Journal of Animal Ecology 56:763–778. Anthony, E.L.P. 1988. Age determination. Pp. 47–57, In T.H. Kunz (Ed.). Ecological and Behavioral Methods for the Study of Bats. Smithsonian Institution Press, Washington, DC. 533 pp. Arnett, E.B. (Ed.) 2005. Relationships between bats and wind turbines in Pennsylvania and West Virginia: An assessment of bat fatality search protocols, patterns of fatality, and behavioral interactions with wind turbines. A final report submitted to the Bats and Wind Energy Cooperative. Bat Conservation International, Austin, TX. 187 pp. Bailey, J.W. 1946. The Mammals of Virginia. Williams Printing Company, Richmond, VA. 416 pp. Barbour, R.W., and W.H. Davis. 1969. Bats of America. The University Press of Kentucky, Lexington, KY. 286 pp. Barclay, R.M.R. 1989a. Long versus short-range foraging strategies of Hoary (Lasiurus cinereus) and Silver-haired (Lasionycteris noctivagans) Bats and the consequence of prey selection. Canadian Journal of Zoology 63:2507–2515. Barclay, R.M.R. 1989b. The effect of reproductive condition on the foraging behavior of female Hoary Bats, Lasiurus cinereus. Behavioral Ecology and Sociobiology 24:31–37. 524 Southeastern Naturalist Vol. 10, No. 3 Blehert, D.S., A.C. Hicks, M. Behr, C.U. Meteyer, B.M. Berlowski-Zier, E.L. Buckles, J.T.H. Coleman, S.R. Darling, A. Gargas, R. Niver, J.C. Okoniewski, R.J. Rudd, and W.B. Stone. 2009. Bat White-Nose Syndrome: An emerging fungal pathogen? Science 323:227. Boyles, J.G., and C.K.R. Willis. 2009. Could localized warm areas inside cold caves reduce mortality of hibernating bats affected by white-nose syndrome? Frontiers in Ecology and the Environment 8:92–98. Brack, V., Jr. 1983. The non-hibernating ecology of bats in Indiana with emphasis on the endangered Indiana Bat (Myotis sodalis). Ph.D dissertation. Purdue University, West Lafayette, IN. 280 pp. Brack, V., Jr. 1985. The foraging ecology of some bats in Indiana. Proceedings from the Indiana Academy of Science 94:231–237. Brack, V., Jr. 2009. Summer bats of Potter and McKean counties, Pennsylvania and adjacent Cattaraugus County, New York. Journal of the Pennsylvania Academy of Science 83(1):17–23. Brack, V., Jr., and J. Duffey. 2006. Bats of Ravenna Training and Logistics Site, Portage and Trumbull counties, Ohio. Ohio Journal of Science 106:130–133. Brack, V., Jr., J.D. Kiser Jr., J. Schwierjohann, and L.B. Williams. 2005. Bats of Camp Dawson, West Virginia: Relative abundance, habitat use, and periods of activity. Proceedings of the West Virginia Academy of Science 77(2):1–6. Braun, E.L. 1950. Deciduous Forests of Eastern North America. Hafner Publishing Company, New York, NY. 596 pp. Broders, H.G., and G.J. Forbes. 2004. Interspecific and intersexual variation in roostsite selection of Northern Long-eared and Little Brown Bats in the Greater Fundy National Park Ecosystem. Journal of Wildlife Management 68:602–610. Carter, T.C., and G.A. Feldhamer. 2005. Roost tree use by maternity colonies of Indiana Bats and Northern Long-eared Bats in southern Illinois. Forest Ecology and Management 219:259–268. Castleberry, S.B., K.V. Miller, and W.M. Ford. 2007. Survey of bat communities in the New River Gorge National River, Gauley River National Recreation Area, and Bluestone National Scenic River: Species occurrence, relative abundance, distribution, and habitat use. Technical Report NPS/NER/NRTR—2007/101. National Park Service. Philadelphia, PA. 42 pp. Cleveland, C.J., M. Betke, P. Federico, J.D. Frank, T.G. Hallam, J. Horn, J.D. Lopez, Jr., G.F. McCracken, R.A. Medellin, A. Moreno-Valdez, C.G. Sansone, J.K. Westbrook, and T.H. Kunz. 2006. Economic value of the pest control service provided by Brazilian Free-tailed Bats in south-central Texas. Frontiers in Ecology and the Environment 4:238–248. Cryan, P.M. 2003. Seasonal distribution of migratory tree bats (Lasiurus and Lasionycteris) in North America. Journal of Mammalogy 84(2):579–593. Dalton, V.M. 1987. Distribution, abundance, and status of bats hibernating in caves in Virginia. Virginia Journal of Science 38(4):369–379. Duchamp, J.E., D.W. Sparks, and R.K. Swihart. 2010. Exploring the “nutrient hot spot” hypothesis at trees used by bats. Journal of Mammalogy 91:48–53. Dzal, Y., L.P. McGuire, N. Veselka, and M.B. Fenton. 2010. Going, going, gone: The impact of white-nose syndrome on the summer activity of the Little Brown Bat (Myotis lucifugus). Biology Letters. Available online at doi:10.1098/rsbl.2010.0859. Accessed 24 November 2010. Fenneman, N.M. 1938. Physiography of the Eastern United States. McGraw-Hill, New York, NY. 714 pp. Fenton, M.B. 1997. Science and the conservation of bats. Journal of Mammalogy 78(1):1–14. 2011 J. Timpone, K.E. Francl, D. Sparks, V. Brack, Jr., and J. Beverly 525 Ford, W.M., M.A. Menzel, and J.A. Menzel. 2002. Influence of summer temperature on sex ratios in Eastern Red Bats (Lasiurus borealis). American Midland Naturalist 147:179–184. Foster, R.W., and A. Kurta. 1999. Roosting ecology of the Northern Bat (Myotis septentrionalis) and comparisons with the endangered Indiana Bat (Myotis sodalis). Journal of Mammalogy 80:659–672. Fujita, M.S., and T.H. Kunz. 1984. Pipistrellus subflavus. Mammalian species 228:1–6. Grindal, S.D., J.L Morissette, and R.M. Brigham. 1999. Concentration of bat activity in riparian habitats over an elevational gradient. Canadian Journal of Zoology 77:972–977. Handley, C.O., Jr. 1979. Mammals of the Dismal Swamp: A historical account. Pp. 297– 357, In P.W. Kirk, Jr. (Ed.). The Great Dismal Swamp. University Press of Virginia. Charlottesville, VA. 427 pp. Handley, C.O., Jr. 1991. Mammals. Pp 539–616. In K. Terwilliger (Ed.). Virginia's Endangered Species: Proceedings of a Symposium. Virginia Department of Game and Inland Fisheries. Blacksburg, VA. 672 pp. Handley, C.O., Jr., G. Tipton, A. Tipton, and D.W. Linzey (Eds.). 1979. Eastern Big- Eared Bat. Pp. 548–549. In Symposium on Endangered and Threatened Plants and Animals of Virginia, Extention Division. Virginia Polytechnic Institute and State University, Blacksburg, VA. 665 pp. Hobson, C.S. 1998. A contribution to the knowledge of the chiropteran fauna of Virginia. M.Sc. Thesis. Department of Biology, Tennessee Technological University, Cookville, TN. 58 pp. Holsinger, J.R. 1964. The Gray Myotis in Virginia. Journal of Mammalogy 45(1):151–152. Hutchinson, J.T., and M.J. Lacki. 1999. Foraging behavior and habitat use of Red Bats in mixed mesophytic forests of the Cumberland Plateau, Kentucky. Pp. 171–177, In J.W. Stringer and D.L. Loftis (Eds.). 12th Central Hardwood Forest Conference. General Technical Report SO-24. US Department of Agriculture, Forest Service, Southern Research Station, Asheville, NC. 303 pp. Johnson, J.B., and J.E. Gates. 2008. Bats of Assateague Island National Seashore, Maryland. American Midland Naturalist 160(1):160–170. Johnson, J.B., J.E. Gates, and M.W. Ford.2008 Distribution and activity of bats at local and landscape scales within a rural-urban gradient. Urban Ecosystems 11:227–242. Johnson, J.B., J.E. Gates, and M.W. Ford. 2009. Notes on foraging activity of female Myotis leibii in Maryland. Research Paper NRS-8. US Department of Agriculture, Forest Service, Northern Research Station. Newton Square, PA. 8 pp. Kunz, T.H. 1982. Lasionycteris noctivagans. Mammalian Species 172:1–5. Kunz, T.H., E.B. Arnett, B.M. Cooper, W.P. Erikson, R.P. Larkin, T. Mabee, M.L. Morrison, M.D. Strickland, and J.M. Szewczak. 2007. Methods and metrics for studying impacts of wind energy development on nocturnal birds and bats. Journal of Wildlife Management 71:2449–2486. Kurta, A. 2010. Reproductive timing, distribution, and sex ratios of tree bats in Lower Michigan. Journal of Mammalogy 91:586–592. Lance, R.F., and R.W. Garrett. 1997. Bat fauna of central Louisiana forests. Texas Journal of Science (Supplement) 49:181–189. Linzey, D.W. 1998. The Mammals of Virginia. McDonald and Woodward Publishing Company, Blacksburg, VA. 459 pp. MacArthur, R.H. 1972. Geographical Ecology. Harper and Row, New York, NY. 299 pp. Menzel, M.A., S.F. Owen, W.M. Ford, B.R. Chapman, K.V. Miller, J.E. Edwards, and P.B. Wood. 2002. Roost tree selection by maternity colonies of Northern Long-eared Bats (Myotis septentrionalis) in an industrial forest of the central Appalachian Mountains. Forest Ecology and Management 155:107–114. 526 Southeastern Naturalist Vol. 10, No. 3 Menzel, M.A., J.M. Menzel, J.C. Kilgo, W.M. Ford, T.C. Carter, and J.W. Edwards. 2003. Bats of the Savannah River Site and vicinity. USDA Forest Service General Technical Report SRS-68. Southern Research Station, Asheville, NC. 69 pp. O'Keefe, J. 2009. Roosting and foraging ecology of forest bats in the southern Appalachian Mountains. Ph.D. Dissertation. Clemson University, Clemson, SC. 102 pp. Owen, S.F., M.A. Menzel, W.M. Ford, J.W. Edwards, J.M. Menzel, B.R. Chapman, P.B. Wood, and K.V. Miller. 2004. Bat activity in harvested and intact forest stands in the Allegheny Mountains. Journal of Applied Forestry 21:154–159. Perry, R.W., and R.E. Thill. 2007. Roost selection by male and female Northern Long-eared Bats in a pine-dominated landscape. Forest Ecology and Management 247:220–226. Racey, P.A. 1988. Reproductive assessment in bats. Pp 31–43, In T.H. Kunz (Ed.). Ecological and Behavioral Methods for the Study of Bats. Smithsonian Institution Press. Washington, DC. 533 pp. Roble, S.M. 2006. Natural Heritage Resources of Virginia: Rare Animal Species. Natural Heritage Technical Report 06-10. Virginia Department of Conservation and Recreation, Division of Natural Heritage, Richmond, VA. 44 pp. Sparks , D.W., and J.R. Choate. 2000. New distribution records of mammals from Kansas. Prairie Naturalist 27:185–192. Stephenson, S. (Ed.). 1993. Upland Forests of West Virginia. McClain Printing, Parsons, WV. 295 pp. Terwilliger, K. (Ed.). 1991. Virginia's Endangered Species: Proceedings of a Symposium. Virginia Department of Game and Inland Fisheries, Blacksburg, VA. 672 pp. Timpone, J.C., J.G. Boyles, K.L. Murray, D.P. Aubrey, and L.W. Robbins. 2010. Overlap in roosting habits of Indiana Bats (Myotis sodalis) and Northern Bats (Myotis septentrionalis). American Midland Naturalist 163:115–123. US Department of Agriculture Forest Service (USDAFS), Southern Region. 2004. Revised land and resource management plan, Jefferson National Forest. Management Bulletin R8-MB 115A. Blacksburg, VA. 396 pp. US Fish and Wildlife Service (USFWS). 1999. Agency draft Indiana Bat (Myotis sodalis) revised recovery plan. Fort Snelling, MN. 53 pp. USFWS. 2007. Indiana Bat (Myotis sodalis) draft recovery plan: First revision. Fort Snelling, MN. 45 pp. Veilleux, J.P., J.O. Whitaker, Jr., and S.L. Veilleux. 2004. Colonies and reproductive patterns of tree-roosting female Eastern Pipistrelles Bats in Indiana. Proceedings of the Indiana Academy of Science 113:60–65. Virginia Department of Game and Inland Fisheries (VDGIF). 1988. Virginia nongame and endangered wildlife investigations. Annual Report: July 1, 1987–June 30, 1988. Blacksburg, VA. 143 pp. VDGIF. 2011. Virginia Fish and Wildlife Information Service. Available online at http:// Webster, W.D., J.F. Parbell, and W.C. Biggs, Jr. 1985. Mammals of the Carolinas, Virginia, and Maryland. University of North Carolina Press, Chapel Hill, NC. 255 pp. Whitaker, J.O., Jr., and V. Brack, Jr. 2002. Myotis sodalis in Indiana. Pp. 53–59, In A. Kurta and J. Kennedy (Eds.). The Indiana Bat: Biology and Management of an Endangered Species. Bat Conservation International, Austin, TX. 253 pp. Whitaker, J.O., Jr., and W.J. Hamilton. 1998. Mammals of the Eastern United States. 3rd Edition. Cornell University Press, Ithaca, NY. 583 pp. 2011 J. Timpone, K.E. Francl, D. Sparks, V. Brack, Jr., and J. Beverly 527 Appendix 1. Bats captured in 11 counties in the Cumberland Plateau and Ridge and Valley physiographic provinces of Virginia, 2000–2009, by gender and age class (adult vs. juvenile; Anthony 1988). Asterisks (*) designate new county records according to the Virginia Department of Game and Inland Fisheries’ Biota of Virginia web-based range. Male Female County Sites Species Total A J A J Bland 27 Big Brown Bat 45 25 2 16 2 Red Bat 12 5 4 1 2 Hoary Bat* 1 0 0 1 0 Eastern Small-footed Bat 2 1 0 1 0 Little Brown Bat 44 13 5 22 4 Northern Bat 90 34 4 44 8 Tri-colored Bat 6 3 0 3 0 Buchanan 6 Big Brown Bat 4 4 0 0 0 Red Bat* 3 3 0 0 0 Silver-Haired Bat* 6 6 0 0 0 Eastern Small-footed Bat* 1 0 0 1 0 Little Brown Bat* 1 1 0 0 0 Northern Bat* 10 2 0 8 0 Dickenson 24 Big Brown Bat 22 14 0 8 0 Red Bat* 23 16 3 4 0 Hoary Bat* 1 1 0 0 0 Eastern Small-footed Bat 7 4 0 3 0 Northern Bat 4 3 0 1 0 Tri-colored Bat* 33 15 0 18 0 Pulaski 6 Big Brown Bat 32 9 2 13 8 Red Bat* 28 4 15 4 5 Hoary Bat* 1 0 0 1 0 Little Brown Bat 12 2 5 4 1 Northern Bat* 19 5 5 4 5 Tri-colored Bat 2 2 0 0 0 Rockingham 16 Big Brown Bat 91 21 18 36 16 Red Bat 37 15 15 2 5 Little Brown Bat 70 25 10 26 9 Northern Bat 11 3 2 5 1 Tri-colored Bat 5 5 0 0 0 Russell 11 Big Brown Bat 7 3 2 2 0 Red Bat* 39 9 17 3 10 Hoary Bat* 1 0 1 0 0 Gray Bat* 5 4 1 0 0 Little Brown Bat 16 3 3 5 5 Tri-colored Bat 10 3 4 1 2 528 Southeastern Naturalist Vol. 10, No. 3 Male Female County Sites Species Total A J A J Shenandoah 12 Big Brown Bat 67 15 10 29 13 Red Bat 39 12 14 2 11 Hoary Bat* 2 0 1 1 0 Little Brown Bat* 43 17 7 16 3 Northern Bat* 16 2 8 3 3 Tri-colored Bat 4 4 0 0 0 Smyth 9 Big Brown Bat 6 0 2 4 0 Little Brown Bat 1 1 0 0 0 Northern Bat* 13 2 0 11 0 Tri-colored Bat 1 1 0 0 0 Tazewell 53 Virginia Big-eared Bat 1 1 0 0 0 Big Brown Bat 109 31 17 55 6 Red Bat* 24 17 3 3 1 Hoary Bat* 1 0 0 1 0 Silver-haired Bat* 7 7 0 0 0 Eastern Small-footed Bat 3 1 1 1 0 Little Brown Bat 47 14 9 18 6 Northern Bat 98 30 12 48 8 Indiana Bat 1 1 0 0 0 Tri-colored Bat 17 17 0 0 0 Wise 35 Big Brown Bat 52 25 8 16 3 Red Bat 70 34 15 5 16 Eastern Small-footed Bat 63 22 9 21 11 Little Brown Bat 7 6 0 0 1 Northern Bat* 42 19 4 13 6 Tri-colored Bat 119 45 50 10 14 Wythe 2 Big Brown Bat 8 0 1 5 2 Tri-colored Bat* 1 1 0 0 0 Total 201 1575 598 289 501 187