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 - john.timpone@hdrinc.com.
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.
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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