United States

Department of

Agriculture

Forest Service

Southern

Research Station

General Technical

Report SRS-68

Bats of the

Savannah River

Site and Vicinity

Michael A. Menzel, Jennifer M. Menzel, John C. Kilgo,

W. Mark Ford, Timothy C. Carter, and John W. Edwards
Authors:

Michael A. Menzel,1 Jennifer M. Menzel,2

John C. Kilgo,3 W. Mark Ford,2 Timothy C. Carter,4

and John W. Edwards5

1

Graduate Research Assistant, Division of Forestry, Wildlife and

Fisheries, West Virginia University, Morgantown, WV 26506;

2

Research Wildlife Biologist, Northeastern Research Station,

USDA Forest Service, Parsons, WV 26287; 3Research Wildlife

Biologist, Southern Research Station, USDA Forest Service, New

Ellenton, SC 29809; 4Graduate Research Assistant, Department of

Zoology, Southern Illinois University, Carbondale, IL 62901; and

5

Assistant Professor, Division of Forestry, Wildlife and Fisheries,

West Virginia University, Morgantown, WV 26506, respectively.

Cover photos:

Clockwise from top left: big brown bats (photo by John

MacGregor); Rafinesque’s big-eared bat (photo by John

MacGregor); eastern red bat (photo by John MacGregor); and

eastern red bat (photo by Julie Roberge).

September 2003

Southern Research Station

P.O. Box 2680

Asheville, NC 28802
i

Bats of the

Savannah River

Site and Vicinity

Michael A. Menzel, Jennifer M. Menzel, John C. Kilgo,

W. Mark Ford, Timothy C. Carter, and John W. Edwards

Abstract

The U.S. Department of Energy’s Savannah River Site supports a diverse

bat community. Nine species occur there regularly, including the eastern

pipistrelle (Pipistrellus subflavus), southeastern myotis (Myotis austroriparius),

evening bat (Nycticeius humeralis), Rafinesque’s big-eared bat (Corynorhinus

rafinesquii), silver-haired bat (Lasionycteris noctivagans), eastern red bat

(Lasiurus borealis), Seminole bat (L. seminolus), hoary bat (L. cinereus), and

big brown bat (Eptesicus fuscus). There are extralimital capture records for

two additional species: little brown bat (M. lucifigus) and northern yellow bat

(Lasiurus intermedius). Acoustical sampling has documented the presence of

Brazilian free-tailed bats (Tadarida brasiliensis), but none has been captured.

Among those species common to the Site, the southeastern myotis and

Rafinesque’s big-eared bat are listed in South Carolina as threatened and

endangered, respectively. The presence of those two species, and a growing

concern for the conservation of forest-dwelling bats, led to extensive and

focused research on the Savannah River Site between 1996 and 2002.

Summarizing this and other bat research, we provide species accounts that

discuss morphology and distribution, roosting and foraging behaviors, home

range characteristics, habitat relations, and reproductive biology. We also

present information on conservation needs and rabies issues; and, finally,

identification keys that may be useful wherever the bat species we describe

are found.

Keywords: Bats, foraging, habitat use, rabies, roosting, Savannah River Site.

A publication of the Savannah River Site National Environmental Research

Park Program. Publication number: SRO-NERP-26
ii
iii

Table of Contents

1 Introduction

2 Major Vegetation Types of

the Savannah River Site

6 The Sitewide Bat Survey

13 Accounts of Species

14 Bats of the Savannah River Site

14 FAMILY VESPERTILIONIDAE

14 Eastern pipistrelle (Pipistrellus subflavus)

Morphology and Distribution .............................................................. 14

Roosting ................................................................................................. 14

Foraging and Home Range .................................................................. 16

Effect of Habitat Type and Stand Age on Flight Activity .................. 16

Reproduction ......................................................................................... 16

18 Southeastern myotis(Myotis austroriparius)

Morphology and Distribution .............................................................. 18

Roosting ................................................................................................. 19

Foraging and Home Range .................................................................. 19

Effect of Habitat Type and Stand Age on Flight Activity .................. 19

Reproduction ......................................................................................... 21

21 Evening bat (Nyticeius humeralis)

Morphology and Distribution .............................................................. 21

Roosting ................................................................................................. 23

Foraging and Home Range .................................................................. 23

Effect of Habitat Type and Stand Age on Flight Activity .................. 23

Reproduction ......................................................................................... 23

25 Rafinesque’s big-eared bat (Corynorhinus rafinesquii)

Morphology and Distribution .............................................................. 25

Roosting ................................................................................................. 26

Foraging and Home Range .................................................................. 26

Effect of Habitat Type and Stand Age on Flight Activity .................. 26

Reproduction ......................................................................................... 26

27 Silver-haired bat (Lasionycteris noctivagans)

Morphology and Distribution .............................................................. 27

Roosting ................................................................................................. 27

Foraging and Home Range .................................................................. 27

Effect of Habitat Type and Stand Age on Flight Activity .................. 28

Reproduction ......................................................................................... 28
iv

Contents, cont.

29 Eastern red bat(Lasiurus borealis)

Morphology and Distribution .............................................................. 29

Roosting ................................................................................................. 29

Foraging and Home Range .................................................................. 30

Effect of Habitat Type and Stand Age on Flight Activity .................. 30

Reproduction ......................................................................................... 32

32 Seminole bat (Lasiurus seminolus)

Morphology and Distribution .............................................................. 32

Roosting ................................................................................................. 33

Foraging and Home Range .................................................................. 34

Effect of Habitat Type and Stand Age on Flight Activity .................. 34

Reproduction ......................................................................................... 34

34 Hoary bat (Lasiurus cinereus)

Morphology and Distribution .............................................................. 34

Roosting ................................................................................................. 35

Foraging and Home Range .................................................................. 35

Effect of Habitat Type and Stand Age on Flight Activity .................. 37

Reproduction ......................................................................................... 37

37 Big brown bat (Eptesicus fuscus)

Morphology and Distribution .............................................................. 37

Roosting ................................................................................................. 39

Foraging and Home Range .................................................................. 39

Effect of Habitat Type and Stand Age on Flight Activity .................. 39

Reproduction ......................................................................................... 39

41 Bats of South Carolina in Areas

Neighboring the Savannah River Site

41 FAMILY VESPERTILIONIDAE

41 Little brown bat (Myotis lucifugus)

Morphology and Distribution .............................................................. 41

Roosting ................................................................................................. 42

Foraging and Home Range .................................................................. 42

Effect of Habitat Type and Stand Age on Flight Activity .................. 42

Reproduction ......................................................................................... 42

43 Eastern small-footed myotis (Myotis leibii)

Morphology and Distribution .............................................................. 43

Roosting ................................................................................................. 44

Foraging and Home Range .................................................................. 44

Effect of Habitat Type and Stand Age on Flight Activity .................. 44

Reproduction ......................................................................................... 44
v

Contents, cont.

44 Northern long-eared myotis (Myotis septentrionalis)

Morphology and Distribution .............................................................. 44

Roosting ................................................................................................. 45

Foraging and Home Range .................................................................. 45

Effect of Stand Age and Habitat Type on Flight Activity .................. 45

Reproduction ......................................................................................... 45

46 Northern yellow bat (Lasiurus intermedius)

Morphology and Distribution .............................................................. 46

Roosting ................................................................................................. 46

Foraging and Home Range .................................................................. 46

Effect of Habitat Type and Stand Age on Flight Activity .................. 47

Reproduction ......................................................................................... 47

48 FAMILY MOLOSSIDAE

48 Brazilian free-tailed bat (Tadarida brasiliensis)

Morphology and Distribution .............................................................. 48

Roosting ................................................................................................. 49

Foraging and Home Range .................................................................. 49

Effect of Habitat Type and Stand Age on Flight Activity .................. 50

Reproduction ......................................................................................... 50

50 Conservation and Research Needs

51 Bats and Rabies

53 Acknowledgments

53 Literature Cited

61 Appendix

61 KEYS TO THE SKINS AND SKULLS

OF SOUTH CAROLINA BATS

Key to the Skins .................................................................................... 65

Key to Skulls.......................................................................................... 69
vi
1

Table 2—Average body mass, wing length, wing-aspect ratio index, and wingloading index of bats of the Savannah River Site

Wing Wing-aspect Wing loading

Species Mass length ratio indexa indexb

g mm

Eastern pipistrelle 5.4 97.7 2.27 1.28

Southeastern myotis 5.7 115.0 2.30 0.99

Little brown bat 6.5 107.7 2.23 1.26

Evening bat 8.0 114.0 2.43 1.49

Rafinesque’s big-eared bat 8.6 127.0 2.27 1.21

Silver-haired bat 9.3 128.3 2.57 1.45

Eastern red bat 10.0 138.0 2.64 1.39

Seminole bat 10.1 138.3 2.66 1.59

Northern yellow bat 20.0 158.3 2.58 2.06

Hoary bat 21.2 183.0 2.94 1.88

Big brown bat 15.3 138.3 2.47 1.97

Brazilian free-tailed batc 11.5 133.0 3.05 2.02

a Wing length/length of fifth phalyx; higher numbers indicate longer, narrower wings.

b [Mass/(wing length x length of fifth phalyx)] x 1,000; higher numbers indicate higher body mass per unit of wing area. c Although the Brazilian free-tailed bat has not been captured on the Savannah River Site, it is likely this species

occasionally occurs onsite.

Table 1—Comparison of average mass, total length, and forearm length of

bats of the Savannah River Site

Species Mass Total length Forearm

g - - - - - - - - mm - - - - - - - -

Eastern pipistrelle 5.4 fg 83.6 h 34.3 g

Southeastern myotis 5.7 efg 86.6 gh 37.1 fg

Little brown bat 6.5 defg 87.4 gh 37.0 fg

Evening bat 8.0 cdefg 86.8 gh 36.2 fg

Rafinesque’s big-eared bat 8.6 cdef 96.1 ef 43.4 cd

Silver-haired bat 9.3 cde 99.5 def 41.6 de

Eastern red bat 10.0 cd 101.8 de 39.5 ef

Seminole bat 10.1 c 103.9 d 41.5 de

Northern yellow bat 20.0 a 127.6 b 51.7 b

Hoary bat 21.2 a 132.5 a 55.2 a

Big brown bat 15.3 b 111.6 c 45.9 c

Brazilian free-tailed bata 11.5 c 93.1 fg 43.4 cd

Means followed by the same letter within a column are not significantly different (P ≤ 0.05). a Although the Brazilian free-tailed bat has not been captured on the Savannah River Site, it is likely this

species occasionally occurs onsite.

Introduction

T 

he Savannah River Site

supports a diverse bat

community. Nine species occur

there regularly. Another three may

use the Site occasionally. These three,

the northern yellow bat (Lasiurus

intermedius Allen), little brown bat

(Myotis lucifugus LeConte), and

Brazilian free-tailed bat (Tadarida

brasiliensis I. Geof. St.-Hilaire), have

been documented on the Site but

probably occur there only occasionally.

Relative to other mammalian orders,

the size of these 12 species varies little.

Total body length ranges from 84 mm

to 132 mm, and wing length ranges

from 98 mm to 183 mm (see tables 1

and 2 and tables accompanying each

species account). In terms of mass,

the largest bat known to occur on the

Site, the hoary bat (Lasiurus cinereus

Beauvois), is only four times larger

than the smallest bat found there,

the eastern pipistrelle (Pipistrellus

subflavus F. Cuvier) (table 1). By

comparison, the largest rodent there,

the beaver (Castor canadensis Kuhl),

is 200 times larger than the smallest

rodent, the eastern harvest mouse

(Reithrodontomys humulis Audubon

and Bachman), and the largest

carnivore, the black bear (Ursus

americanus Palla), is more than

900 times larger than the smallest

carnivore, the long-tailed weasel

(Mustela frenata Lichtenstein)

(Cothran and others 1991).

Distributions of most temperate

zone bats are strongly influenced

by the availability of roost structures

(Findley 1993, Humphrey 1975).

The roosting habits of bats in South

Carolina are diverse (table 3), resulting

in variations in the diversity of bat

communities across the State. The

least diverse bat community occurs in

the Piedmont (6 species), whereas the

most diverse occur in the Blue Ridge

and lower Coastal Plain (12 species)

(Menzel and others 2003). Nine of the

fourteen species that occur in South

Carolina (Menzel and others 2003)

typically roost in caves or mines during
2 Bats of the Savannah River Site and Vicinity

part of the year (table 3). Three

species roost in mines or caves, but

only during the winter (table 3).

In addition to suitable foliage and

cavity roost sites, the Blue Ridge also

contains mines and crevices in vertical

rock faces, resulting in the greatest

diversity of roost structures in the

State. Thus diversity in the bat

community on the Savannah River

Site may be somewhat limited simply

by its physiographic position.

In the last 10 years many studies

have focused on the distribution

and natural history of bats in the

Southeastern United States. Over

half the total published literature

about the natural history of bats in

South Carolina and Georgia was

published in that period, and research

has proceeded as fast in other

Southeastern States. Although many

aspects of their behavior and natural

history in the Southeast remain poorly

understood, much is known about

bat species’ roosting requirements,

foraging behavior, diet, home range,

and reproduction. Making land

management decisions without regard

for the effects they may have on bat

communities no longer can be justified

by claims that too little is known about

habitat requirements of this diverse

mammalian order.

Many recent studies of bat habitat

requirements were conducted on the

Site. Similar studies elsewhere in

the Southeast also provided useful

information about the natural history

of bats known to occur there. This

report summarizes information about

the distribution, roosting ecology,

foraging behavior, diet, home range,

and reproduction of bats that occur

on and near the Site. Information

contained herein should aid land

managers in assessing how their

management decisions may affect

bat species.

Major Vegetation

Types of the

Savannah River Site

T he Savannah River Site is

a former nuclear weapons

material production facility

operated by the U.S. Department

of Energy (DOE). It borders the

Savannah River in west-central South

Carolina in the upper Coastal Plain

physiographic province and occupies

78 000 ha (fig. 1). It is located in parts

of Aiken, Barnwell, and Allendale

Counties. When the DOE acquired the

land in 1950, about 33 percent of it was

cropland or pasture, and the remainder

was forested (see Workman and

McLeod 1990 for a detailed description

of vegetation communities). In 1952

the USDA Forest Service began

managing timber on the Site, and 20

years later it was designated the

Nation’s first National Environmental

Research Park (Workman and McLeod

1990). The Site has a temperate climate

characterized by mild winters and

long summers with average air

temperatures of 9 and 27 o

C,

respectively (Workman and McLeod

1990). The frost-free period lasts

roughly 240 days, and the average

annual rainfall is 120 cm. Most rain

falls in March (13.1 cm) and the least

in November (5.9 cm) (Workman and

McLeod 1990).

Using Workman and McLeod (1990)

and Imm and McLeod (in press),

we identified seven major vegetation

types on the Site (fig. 2). These include

loblolly-slash pine (Pinus taeda L., P.

elliottii Engelm.) (32 461 ha), longleaf

pine (P. palustris Mill.) (17 008 ha),

mixed pine-hardwood (4569 ha),

upland hardwood (2738 ha), and

bottomland hardwood (16 462 ha)

forests; lakes/ponds/marshes (1672

ha), and grass-brush (5214 ha).

Pine forests dominate the Savannah

River Site landscape, and pine canopies

there tend to be dominated by a single

species. Because of their age and

management on the Site, longleaf pine

stands tend to have lower densities and

Table 3—Typical roosting habitats of the 14 species of bats that occur in South Carolina

Bark/ Artificial Cave/ Rock

Species Foliage cavity structure mine crevices

Eastern pipistrelle S S S WS

Southeastern myotis S S WS

Little brown bat S S WS

Small-footed myotisa S WS

Northern long-eared myotisa S WS W

Evening bat S S S

Rafinesque’s big-eared bat WS WS WS

Silver-haired bat WS W W S

Eastern red bat WS

Seminole bat WS

Northern yellow bat WS

Hoary bat WS

Big brown bat WS WS WS

Brazilian free-tailed bata WS WS WS

W = winter roost; S = summer roost.

a Species that have not been captured on Savannah River Site.
3

Figure 1—Location of South Carolina’s four physiographic provinces and the Savannah River Site.

1 - Blue Ridge

2 - Piedmont

3 - Upper Coastal Plain

3 - Lower Coastal Plain

turkey oak (Quercus laevis Walt.),

scrub post oak (Q. margaretta Sarg.),

bluejack oak (Q. incana Bartr.),

blackjack oak (Q. marilandica

Muenchh.), blackgum (Nyssa sylvatica

Marsh.), sand hickory [Carya pallida

(Ashe) Engl. & Graebn.], mockernut

hickory [C. tomentosa (Poir.) Nutt.],

sassafras [Sassafras albidum (Nutt.)

Nees], and sparkleberry (Vaccinium

arboreum Marsh.) (Workman and

McLeod 1990). On more mesic upland

and bottomland sites, loblolly pine

forms mixed stands with the hardwood

species typical of such sites.

Upland hardwood communities on

the Site are dominated by oaks and

hickories and are located on the flats of

the Aiken Plateau, on stream bluffs,

basal areas than loblolly and slash pine

forests. Because vegetation density is

a primary factor affecting bat foraging

activity (Owen and others, in press),

we treated longleaf forest separately

(fig. 2). Both forest types are managed

for timber production on the Site

within the constraints of red-cockaded

woodpecker (Picoides borealis Vieillot)

habitat management guidelines.1

See Blake (in press) for a detailed

discussion of forest management

practices there. Although the rotation

length for pines there is relatively long

for managed forests in the Southeast

(50 to 120 years), many age classes

are found in the region. Clearcut

harvesting is used to convert slash

pine to species better suited for sitespecific conditions, i.e., longleaf,

loblolly pine. Young pine plantations

constitute most early successional

habitat on the Site. Managers there

use prescribed fire on a 3- to 5-year

rotation to treat areas of pine forest.

Mixed pine-hardwood forests occupy

much of the Savannah River Site,

ranging from the dry and infertile

sandhills, which are remnant beach

dunes from the Cretaceous shoreline,

to moist, rich bottomland sites, which

are subject to periodic inundation.

Tree species occurring in sandhill

communities include longleaf pine,

1

 Edwards, J.W.; Smathers, W.M., Jr.; Lemaster,

E.T.; Jarvis, W.L. 2000. Savannah River Site redcockaded woodpecker management plan. 66 p.

On file with: USDA Forest Service-Savannah

River, P.O. Box 700, New Ellenton, SC 29809.

Major Vegetation Types of the Savannah River Site
4 Bats of the Savannah River Site and Vicinity

and along the borders

of bottomland forests

(Workman and McLeod

1990). Upland forests

typically contain mixtures of

white oak (Q. alba L.), post oak

(Q. stellata Wangenh.), black oak

(Q. velutina Lam.), scarlet oak (Q.

coccinea Muenchh.), southern red

oak (Q. falcata Michx.), blackjack

oak, laurel oak (Q. laurifolia Michx.),

mockernut hickory, sweetgum

(Liquidambar styraciflua L.),

dogwood (Cornus florida L.),

sassafras, and Georgia hackberry

(Celtis occidentalis L. var. georgiana).

In addition, stream-bluff upland

hardwood communities may contain

tulip poplar (Liriodendron tulipifera

L.), beech (Fagus grandifolia Ehrh.),

red buckeye (Aesculus pavia L.),

sourwood [Oxydendrum arboreum

(L.) DC.], and witch-hazel

(Hamamelis virginiana L.). Upland

hardwood communities are denoted

as upland hardwoods on the forest

habitat maps included in each species

account (fig. 2).

Bottomland hardwood forests on

the Site are typical of the mesic

mixed-hardwood forests of the

southeastern Coastal Plain (Workman

and McLeod 1990). Bottomland

hardwood communities occur in the

lowlands at elevations between 25

and 90 m and commonly are flooded

during high-water periods in late

winter and early spring (Workman

and McLeod 1990). Most hardwoods

on the Site were selectively harvested

in the late 1800s and early 1900s.

They remained relatively undisturbed

Figure 2—The 449 locations where AnaBat surveys were conducted during summer

2001, and the distribution of 7 habitat types across the Savannah River Site.

during the last three-quarters of the

20th century (Workman and McLeod

1990). Common tree species in the

bottomland forests include sweetgum,

red maple (Acer rubrum L.), swamp

chestnut oak (Q. michauxii Nutt.),

cherrybark oak (Q. falcata var.

pagodifolia Ell.), water oak (Q. nigra

L.), laurel oak, willow oak (Q. phellos

L.), overcup oak (Q. lyrata Walt.),

green ash (Fraxinus pennsylvanica

Marsh.), and blackgum. Because the

structure of most swamp forests on

the Site—occurring primarily along

the Savannah River (Whipple and

others 1981)—is similar to the

structure found in bottomland

hardwood forests, we displayed

them as bottomland hardwoods

on the forest habitat maps included

with each species (fig. 2), and we

otherwise included them with

bottomland hardwood communities.

Swamp forests occur at elevations

between 25 and 35 m and commonly

are inundated during a portion of the

year (Workman and McLeod 1990).

Approximately half of the swamp

4
5

Figure 3—Location of streams and Carolina bays on the Savannah River Site.

Carolina bays

Water bodies

forests on the Site are second-growth

bald cypress [Taxodium distichum

(L.) Rich. var. distichum])-water

tupelo (N. aquatica L.) swamp, and

the other half is composed of ridges

and hardwood islands (Jensen and

others 1984). Common tree species

in swamp forest areas include bald

cypress, water tupelo, water ash

(F. caroliniana Mill.), green ash,

American elm (Ulmus americana

L.), and sycamore (Platanus

occidentalis L.).

Most of the acreage in lakes,

ponds, and marshes is in the

reservoirs of Par Pond and L

Lake, which the DOE

constructed for use in

cooling reactor. Although

not extensive, an important

habitat included in this category

is Carolina bays. Carolina bays

are natural shallow elliptically shaped

wetlands whose depths range from

zero to a few meters (Schalles and

others 1989). Although they occur

from Florida to Virginia, most

Carolina bays are in North and

South Carolina. Typically they are

oriented northwest to southeast.

Approximately 500,000 Carolina bays

occur on the Coastal Plain, and about

300 are on the Savannah River Site

(Kirkman and others 1996). Most

Carolina bays are surrounded by

upland pine-mixed hardwood forest.

Bat activity commonly is concentrated

over the wetland habitat that Carolina

bays provide, because overstory

vegetation is sparse and aquatic

insects congregate around them

(Menzel and others 2003). Because

Carolina bays are small and occur

in patchy distribution across the

landscape, it is hard to map the

patterns of bat activity relative to

them (see figure 3 for the location of

Carolina bays). To simplify the foresthabitat maps included with each

species account, we did not identify

Carolina bays, although we did

include them in the figures that

compare bat activity levels among

the habitat types.

Grass-brush habitats include roads,

railroad and utility rights-of-way, and

lawns around the Site’s facilities areas.

The facilities themselves are shown

on the vegetation maps. Such grassy

areas constitute the only permanently

maintained early successional habitat

on the Site. Typically, the rights-of-way

are dominated by a mix of broomsedge

(Andropogon spp.), blackberry (Rubus

spp.), Japanese honeysuckle

(Lonicera japonica), and

various grasses.

Because of their importance

to foraging bats, we delineated

separately the sampling points located

within canopy gaps (regardless of

vegetation type) and included them

on maps displaying the Carolina bays

and seven vegetation types.

Major Vegetation Types of the Savannah River Site
6 Bats of the Savannah River Site and Vicinity

The Sitewide Bat Survey I

n preparing the species accounts,

we relied heavily on a survey

conducted during summer 2001.

In that survey we sampled 449 points

across the Site, covering areas

representative of most site habitats

(fig. 2). We conducted the surveys

between sunset and 02:30 from late

May to early August 2001 [see Menzel

(2003) for a detailed description of the

methods]. At each sampling point, we

censused bat activity for 20 minutes

using an AnaBat II bat detection

system (Titley Electronics, Ltd.,

Ballina, New South Wales, Australia)

(Murray and others 2001). We

identified calls using both qualitative

and quantitative techniques (Menzel

1998, O’Farrell and others 1999) and

a call library, which we created by

recording the calls of hand-released

bats captured in the Southeast. Figures

4 and 5 are illustrations generated

using AnaLook software (4.8i), and

they represent typical search phase

calls of the nine bat species occurring

regularly on or around the Site.

Menzel (2003) was not able to

discriminate between the calls of

eastern red (Lasiurus borealis) and

Seminole (L. seminolus) bats based on

the calls recorded, so both species are

included in a group termed “eastern

red/Seminole bat.” We determined the

age and vegetation type of the stand

around each survey point using 1985

Forest Inventory and Analysis data

provided by the USDA Forest Service.

The following results are condensed

from Menzel (2003). Trends in bat

activity were relatively consistent

among habitat types. The activity level

(number of calls per 20-minute survey)

of the eastern red/Seminole bat group

was highest, followed by eastern

pipistrelles, evening bats, big brown

and hoary bats, and southeastern

myotis (in order of decreasing activity)

(fig. 6). Because of the highly variable

nature of bat activity across the Site, we

found few differences in the levels of

activity among species within each

habitat type (table 4). Overall, we

Table 4—Comparison of flight activity (number of calls per 20-minute survey) among 6 bat species in 10 habitat types on the Savannah River Site during summer 2001

Eastern red/ Eastern Southeastern

Survey Seminole bats pipistrelles Evening bats Big brown bats Hoary bats myotis

Habitat type locations Mean SE Mean SE Mean SE Mean SE Mean SE Mean SE F P

no.

Lakes and ponds 16 3.3A 1.11 2.7AB 1.45 0.6BC 0.24 0.8BC 0.47 0.4BC 0.26 0.1C 0.09 2.93 0.017Carolina bays 35 6.1A 2.25 0.8B 0.23 1.7B 0.61 0.5B 0.25 0.1B 0.06 0.4B 0.15 5.64 0.001Bottomland hardwoods 66 4.0A 0.94 2.0AB 0.97 1.2B 0.50 0.6B 0.28 1.3B 1.20 0.4B 0.15 2.89 0.014Forest gaps 38 3.5A 1.11 1.1B 0.56 0.6B 0.27 0.1B 0.09 0.0B 0.03 0.8B 0.29 5.74 0.001Grass-brush 59 5.7A 1.69 2.1B 0.71 2.4B 1.05 0.8B 0.42 0.1B 0.07 0.1B 0.06 5.73 0.001Upland hardwoods 11 2.0A 1.03 0.5B 0.45 0.5B 0.31 0.0B 0. 0.0B 0. 0.1B 0.09 2.56 0.036Pine hardwoods 14 4.4A 1.72 0.6B 0.34 0.1B 0.10 0.1B 0.07 0.0B 0. 0.1B 0.14 5.83 0.001Loblolly-slash pines 140 2.0A 0.41 0.5B 0.20 0.2B 0.06 0.1B 0.03 0.1B 0.03 0.4B 0.17 13.22 0.001Longleaf pines 70 3.0A 0.96 0.3B 0.23 0.6B 0.17 0.2B 0.11 0.0B 0.03 0.1B 0.07 7.75 0.001

Means followed by the same letter within a column are not significantly different (P ≤ 0.05).
7

Figure 4—Illustrations generated using AnaLook software (4.8i) of typical search-phase calls of four low-frequency bat species thatoccur on or around the Savannah River Site: (A) big brown bat, (B) hoary bat, (C) silver-haired bat, and (D) Brazilian free-tailed bat.

The Sitewide Bat Survey
8 Bats of the Savannah River Site and Vicinity

Figure 5—Illustrations generated using AnaLook software (4.8i) of typical search-phase calls of four high-frequency bat species that occur on or around

the Savannah River Site: (A) eastern red/Seminole bat, (B) evening bat, (C) eastern pipistrelles, and (D) southeastern myotis.
9

9876543210

Lakes and ponds

Carolina bays

Bottomland hardwoods

Forest gaps

Grass-brush

Upland hardwoods

Pine-hardwoods

Loblolly-slash pines

Longleaf pines

Habitat types

Mean number calls per 20 minute survey

Eastern red/Seminole bat

Eastern pipistrelleEvening batBig brown batHoary bat

Southeastern myotis

The Sitewide Bat Survey Figure 6—Effect of habitat type on the flight activity of six bat species on the Savannah River Site. ‘
10 Bats of the Savannah River Site and Vicinity

0

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

Lake/pond

Carolina bays

Bottomland hardwood

Forest gaps

Grass/brush

Upland hardwood

Pine-hardwood

Loblolly-slash pine

Longleaf pine

Habitat types

Mean number of calls per 

20 minute survey

A

B

Figure 7—(A) AnaBat survey locations where bat activity was detected in surveys conducted during summer 2001

and (B) a comparison of bat flight activity among vegetation community types on the Savannah River Site.
11

Table 5—Comparison of flight activity (number of calls per 20-minute survey) among six bat species in five stand-age classes on the Savannah River Site

during summer 2001

Eastern red/ Eastern Southeastern

Survey Seminole bats pipistrelles Evening bats Big brown bats Hoary bats myotis

Age class locations Mean SE Mean SE Mean SE Mean SE Mean SE Mean SE F P

no.

Clearcut 6 5.3 2.55 3.3 3.33 2.0 0.93 0.8 0.65 0. 0. 0.2 0.17 5.30 0.0014–20 135 3.7A 0.84 1.3B 0.36 1.2B 0.47 0.5B 0.20 0.1B 0.04 0.3B 0.12 9.61 0.00121–40 36 3.0A 1.04 0.4B 0.22 0.4B 0.21 0.1B 0.07 0.0B 0.03 1.0B 0.49 22.27 0.00141–60 190 3.5A 0.63 0.5B 0.17 0.6B 0.14 0.2B 0.06 0.0B 0.02 0.2B 0.07 2.47 0.032> 60 66 3.4A 0.70 1.7AB 0.94 1.1B 0.50 0.4B 0.24 1.3B 1.20 0.3B 0.15 1.37 0.264

Means followed by the same letter within a column are not significantly different (P ≤ 0.05).

detected bat activity on 87.5 percent of

the lake and pond survey points, 80.0

percent of the Carolina bay survey

points, 66.7 percent of the bottomland

survey points, 71.1 percent of the

forest gap survey locations, 74.6

percent of the grass-brush survey

points, 63.6 percent of the upland

hardwood points, 57.1 percent of the

pine-hardwood points, 48.6 percent of

the loblolly-slash survey points, and

58.6 percent of the longleaf survey

locations. Sitewide, we recorded bat

activity at 62.6 percent of all survey

locations (fig. 7). For all species

combined, activity was highest in

grass-brush and bottomland

hardwoods and over lakes, ponds,

and Carolina bays. Levels of activity

tended to be low in upland hardwood

and pine-hardwood and on sites with

pine-dominated canopies (fig. 7).

In all forest age classes except

clearcut, bat activity levels also

differed among the species (table 5).

This trend was consistent among age

classes other than clearcut. The level

of activity of the eastern red/Seminole

bat group was highest, followed by

eastern pipistrelles, evening bats,

big brown and hoary bats, and

southeastern myotis (table 5). For all

species combined, activity was highest

in clearcuts, moderate in stands aged 4

to 20 years and > 60 years, and lowest

in stands aged 21 to 60 years (fig. 8).

This trend corresponds with the

typical density of vegetation in stands

in these age classes. In clearcuts, bat

foraging activity is not impeded by

forest clutter. The area above newly

regenerating stands (4 to 20 years)

provides excellent foraging habitat.

Stands aged 21 to 60 years often are

stocked so densely that there is little

room for bats to forage. After stands

reach > 60 years, their canopies begin

to open—either through natural stand

attrition or as a result of thinning

treatments—and these openings

provide foraging areas.

The Sitewide Bat Survey
12 Bats of the Savannah River Site and Vicinity

Figure 8—Effect of stand age on bat flight activity on the Savannah River Site.

0

2

4

6

8

10

12

14

16

18

Clearcut 4-20 21-40 41-60 >60

Age class (year)

Mean number calls per 20 minute survey

0

1

2

3

4

5

6

7

8

Clearcut 4-20 21-40 41-60 >60

Age class (year)

Mean number calls per 20 minute survey

Eastern red/Seminole bats

Eastern pipistrelle

Evening bat

Big brown bat

Hoary bat

Southeastern myotis

Figure 9—Effect of stand age on the flight activity of six bat species on the Savannah River Site.
13

The foraging activity patterns of two

species did not correspond to the

activity pattern of all species combined

(fig. 9). Hoary bat activity was highest

in stands aged > 60 years, and no

hoary bat calls were detected in

clearcuts (fig. 9). Southeastern myotis

activity was highest in stands 21 to 40

years old and lowest in the youngest

and oldest stands.

Accounts of Species

Species accounts are organized

into two major sections, “Bats

of the Savannah River Site” and

“Bats of South Carolina in Areas

Neighboring the Savannah River

Site.” Twelve bat species have been

documented on the Site, but not all

occur there regularly. To determine

those species that do, we examined the

range of each of the 14 bat species

found in South Carolina (Menzel and

others 2003), reviewed museum

records for specimens captured on

the Site itself, and reviewed capture

records maintained for bat surveys

taken there. In summarizing previous

capture and collection records, we

reviewed museum records listed

in Menzel and others (2003) for

specimens captured on the Site;

recorded unpublished survey records

from 1999 to 2001 for one of the

authors;2

 recorded capture records

reported by Carter (1998), Menzel

(1998), and Childs and Buchler (cited

in Cothran and others 1991); reviewed

records maintained by public health

personnel in South Carolina, as

reported by DiSalvo and others

(2002); and reviewed records

maintained by the South Carolina

Department of Natural Resources.

Based on our review, some species

captured on the Site probably do

not occur there regularly and should

be of little concern to Site managers,

e.g., the northern yellow bat (L.

intermedius) and the little brown

bat (M. lucifugus). In addition,

although the Brazilian free-tailed bat

(T. brasiliensis) has not been captured,

it has been detected on call surveys,

so it occurs occasionally.

Within each species account, we

tried to synthesize all information

collected in South Carolina about

roosting ecology, foraging behavior,

diet, home range, and reproduction.

We used information from published

studies, faunal surveys of specific

regions within the State (Cothran and

others 1991, Golley 1966, Penney 1950,

Sanders 1978), unpublished master’s

theses (Carter 1998, Menzel 1998),

and unpublished museum records.

Because aspects of the natural history

of some bat species found on the Site

have not been studied anywhere in

South Carolina, we supplemented

information in each species account

with information collected from other

regions. Where information collected

from other regions was used, we noted

it in the individual species accounts.

In each species account, we

included the following subsections—

Morphology and Distribution,

Roosting, Foraging and Home Range,

Effect of Stand Age and Habitat Type

on Flight Activity, and Reproduction.

Morphology and Distribution

describes the species’ general physical

characteristics, subspecies that occur

on the Site, and species distribution.

That section includes a picture of the

species and a table providing its

average body measurements. Data

provided in the body measurement

table were collected from specimens

captured or collected in the Southeast

(primarily Georgia). If the average

body measurements differed

significantly (t-test, alpha < 0.05)

between males and females, we

provided a separate table for each sex.

Each Morphology and Distribution

section references a map illustrating

distribution of the species in South

Carolina, adapted from Menzel and

others (2003).

The Roosting section includes

information about a species’ summer

and winter roosting habits.

Information about the general roosting

habits of each species also is provided.

In addition, specific information

concerning roosting habits in South

Carolina or on the Site is provided for

seven of the nine bat species that

regularly occur on-site.

The Foraging and Home Range

section includes information about

emergence time, foraging habitats,

specific foraging characteristics, e.g.,

foraging height, whether the species

has ever been documented foraging in

groups, and diet. When data were

available, we included information

about selective foraging. Also where

available, we included information

about the spatial attributes and size of

the species’ home range. We defined

home range as the spatial extent an

individual traveled during 3 to 15

nights. All reported home ranges were

based on > 25 telemetry points and

most were based on > 30 points.

In each species account, we discuss

habitat types on the Site in which a

species’ foraging activity was

concentrated and the habitat types it

seemed to use less. This information is

based largely on the sitewide surveys

conducted during summer 2001. We

used survey data to prepare maps

showing the general distribution of

each species’ foraging activity across

the Site. We also prepared figures for

each species or species group, i.e.,

eastern red/Seminole bat, that show

the average level of call activity in each

vegetation type.

We conclude each account with

a brief discussion of the species’

reproductive habits. Because little

work has been done with regard

to bat reproductive habits in South

Carolina, most of this information

came from studies conducted in

other regions of the United States.

Information included in this section

includes the time of year mating

occurs, whether delayed fertilization

occurs, when the young are born, and

the average number of young per litter.

2

 Menzel, M.A. Unpublished data. On file with:

West Virginia University, Division of Forestry,

Percival Hall, Morgantown, WV 26506.

Accounts of Species
14 Bats of the Savannah River Site and Vicinity

Bats of the

Savannah River Site

FAMILY VESPERTILIONIDAE

Vespertilionidae is the largest

family of bats, consisting of

42 genera and 355 species

(Nowak 1994). Family distribution

is cosmopolitan, and members are

found on every continent except

Antarctica. Its members are found

in habitats ranging from desert to

tropical forest. Because members of

this group lack noseleafs and have

simple, unmodified lips and nostrils,

they are commonly called the plainfaced bats. The tragus usually is well

developed, and the tail is not free from

the uropatagium.

Nine genera and thirty-one species

of vespertilionids occur in the United

States. Of these, 8 genera and 13

species occur in South Carolina.

All nine bat species that commonly

occur on the Savannah River Site are

members of the family Vespertilionidae.

Eastern pipistrelle

(Pipistrellus subflavus)

Morphology and Distribution

The eastern pipistrelle is the second

smallest bat found in South Carolina

and the smallest bat on the Site

(fig. 10). One of four recognized

subspecies, P. s. subflavus, occurs in

the State (Fujita and Kunz 1984). Mass

ranges from 3.3 to 8.0 g, and average

total length is 85.1 mm (Whitaker and

Hamilton 1998). See table 6 for average

body measurements of individuals

collected in the Southeast.

The eastern pipistrelle is common

throughout much of the eastern half

of North and Central America. In the

United States, its range extends west

from Maine to Minnesota and south

through much of east Texas (Fujita

and Kunz 1984). Thirty-nine eastern

pipistrelles from South Carolina are

preserved in collections, and 23 have

been live-captured in the State (Menzel

and others 2003). The eastern

pipistrelle occurs in all of South

Carolina’s physiographic provinces

and is common throughout the State

(fig. 11) (Menzel and others 2003).

Eastern pipistrelles were captured

in late June and late August during

survey work conducted by Childs and

Buchler in 1979 (Anon. 1980, Cothran

and others 1991). Between 1996 and

2001, we captured one female and five

male eastern pipistrelles on the Site

(table 7). Based on the regional

distribution of this species, it is likely

that while it is not abundant on the

Site, the eastern pipistrelle is a

common resident there.

Roosting

Eastern pipistrelle hibernacula have

been found in culverts (Moore 1949),

storm sewers (Goehring 1954), tunnels

(Mohr 1942), caves (Davis 1966, Hahn

Figure 10—Eastern pipistrelle.

Photo courtesy of John MacGregor
15

Figure 11—Distribution of the eastern pipistrelle in South Carolina (Menzel

and others 2003). Symbols denote counties for which records exist.

Eastern pipistrelle

(Pipistrellus subflavus)

Table 6—Body measurements of the eastern pipistrelle bat in the Southeastern

United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - - - - - mm - - - - - - - - - - - - - - - - - -

Female

Total 15 86.3 5.96 6.90 77.0 – 99.0

Tail 15 39.0 3.16 8.11 34.0 – 45.0

Foot 15 7.8 1.21 15.48 6.0 – 10.0

Ear 15 12.8 1.56 12.16 10.0 – 14.5

Tragus 14 4.7 1.37 28.98 3.5 – 7.5

Forearm 15 34.6 1.12 3.24 33.0 – 36.0

- - - - - - - - - - - - - - - - - - - g - - - - - - - - - - - - - - - - - - -

Mass 14 5.7 1.03 18.06 4.3 – 8.0

- - - - - - - - - - - - - - - - - - mm - - - - - - - - - - - - - - - - - -

Male

Total 26 82.1 5.74 6.99 71.0 – 95.0

Tail 26 36.9 2.95 7.98 30.0 – 41.0

Foot 26 8.3 1.28 15.47 6.5 – 11.0

Ear 26 12.5 1.09 8.71 10.5 – 14.0

Tragus 22 4.7 1.48 31.12 3.0 – 8.0

Forearm 24 34.2 1.96 5.75 30.0 – 39.0

- - - - - - - - - - - - - - - - - - - g - - - - - - - - - - - - - - - - - - -

Mass 24 5.2 1.16 22.50 3.3 – 8.0

Measures that differed significantly between the sexes (P ≤ 0.05). a Number of individuals measured.

Bats of the Savannah River Site

1908, Raesly and Gates 1987, Swanson

and Evans 1936), and mines (Menzel

and others 1997, Sealander and Heidt

1990, Whitaker and Rissler 1992). No

studies have quantitatively examined

its winter roosting habits in South

Carolina, although Golley (1966)

collected a male eastern pipistrelle

roosting under a house in Berkeley

County, SC, during the winter. Golley

(1966) also found a large colony of

eastern pipistrelles roosting in caves

near Parler in Orangeburg County,

SC, during the winter and spring.

The winter roosting habits of this

species on the Savannah River Site

are unknown.

The eastern pipistrelle’s summer

roosting habits also are poorly known.

Maternity roosts usually are found in

human-made structures such as barns

(Lane 1946, Poole 1938) or houses

(Allen 1921). Menzel and others (1996)

captured a female and her nonvolant

young in a pitfall trap set near a basal

cavity in a large sweetgum, which

suggests that basal cavities may serve

as maternity roosts for the species. A

few individuals also have been found

roosting in tree canopies (Findley

1954) and Spanish moss (Tillandsia

usneoides) during the summer

(Jennings 1958, Menzel and others

1999). In summer 1997, Carter and

others (1999b) located an eastern

pipistrelle roosting in swamp chestnut

oak, sweetgum, and laurel oak in a

bottomland hardwood stand on

the Site.
16 Bats of the Savannah River Site and Vicinity

Table 7—Number of bats captured on the Savannah River Site,

1996 to 2000

Age

Species Sex Adults Juveniles

Eastern pipistrelle (n = 7) F 1 0

M 50

Southeastern myotis (n = 2) F 0 0

M 10

Little brown bat (n = 1) F 1 0

M 00

Evening bat (n = 67) F 27 13

M 21 6

Rafinesque’s big-eared bat (n = 9) F 7 0

M 10

Eastern red bat (n = 56) F 30 11

M 85

Seminole bat (n = 22) F 6 6

M 73

Hoary bat (n = 3) F 2 0

M 00

Big brown bat (n = 6) F 2 1

M 30

Total (n = 173) F 70 31

M 52 14

n = total number captured (the difference between n and sum of age and sex data is

because age and sex were not recorded for all individuals captured);

F = female; M = male.

Foraging and Home Range

Eastern pipistrelles commonly

forage over waterways and along

field edges (Fujita and Kunz 1984,

Whitaker and Hamilton 1998). Harper

(1927) found them foraging over fields

and cypress bays and in pine barrens

of the Okefenokee Swamp. On the

Savannah River Site, Carter and others

(1999b) documented foraging among

bottomland hardwoods and in pine

stands. Their diet includes insects in

the orders Coleoptera, Homoptera,

Diptera, Hymenoptera, and Lepidoptera

(Ross 1961, Sherman 1939, Whitaker

1972). They also forage on

trichopterans and hemipterans

(Carter and others 1999b).

Little is known about this species’

home range, although Krishon and

others (1997) documented the home

range of one eastern pipistrelle on

Sapelo Island, GA. It was 389.2 ha and

concentrated in high marsh (47

percent), oak (24 percent), and slashloblolly pine (17 percent) habitats. The

average distance from the roost area to

a foraging location was 1137 m. Carter

and others (1999b) documented an

eastern pipistrelle’s home range on

the Site at 395.5 ha, concentrated in

bottomland hardwood and upland

pine stands.

Effect of Habitat Type and Stand

Age on Flight Activity

We recorded eastern pipistrelles at

37.5 percent of the lake and pond

survey points, 28.6 percent of the

Carolina bay survey points, 18.2

percent of the bottomland hardwood

survey points, 18.4 percent of the

forest gap survey locations, 23.7

percent of the grass-brush survey

points, 9.1 percent of the upland

hardwood points, 21.4 percent of the

pine-hardwood points, 12.1 percent of

the loblolly-slash survey points, and

7.1 percent of the longleaf survey

locations. Overall, we recorded the

eastern pipistrelle’s call at 16.7 percent

of the survey locations (fig. 12).

Although variable, its activity was

greatest around lakes and ponds,

bottomland hardwood forests, and

grass-brush (fig. 12). In bottomland

hardwoods, we recorded a call

approximately every 10 minutes. The

figures comparing bat activity among

vegetation community types are

arranged with hydric communities on

the left, mesic communities in the

center, and xeric communities on the

right. Eastern pipistrelle activity was

concentrated in hydric and mesic

communities (table 4).

Eastern pipistrelle activity also

differed among stands of different

ages (fig. 9). We recorded most in

clearcuts (fig. 9) or areas devoid of

vegetation, e.g., roads, open water

habitats (fig .12). We recorded

moderate levels of activity in stands

aged 4 to 20 years and > 60 years (fig.

9). The species’ activity was low in

stands 21 to 60 years old (table 5).

Reproduction

Eastern pipistrelles mate in autumn

and spring (LaVal and LaVal 1980) and

parturition occurs in May and June

(Jennings 1958, LaVal and LaVal 1980).

The sexes segregate into bachelor and

maternity colonies in the spring. No

information is available concerning the

parturition times for this species in

South Carolina, but on the Savannah

River Site it probably occurs in May

and June.
17

0.0

0.5

1.0

1.5

2.0

2.5

3.0

3.5

4.0

4.5

Lake/pond

Carolina bays

Bottomland hardwood

Forest gaps

Grass/brush

Upland hardwood

Pine-hardwood

Loblolly-slash pine

Longleaf pine

Habitat types

Mean number of calls per

20 minute survey

A

B

Figure 12—(A) AnaBat survey locations where eastern pipistrelles were detected in surveys conducted during summer 2001 and

(B) a comparison of flight activity levels of eastern pipistrelles among vegetation community types on the Savannah River Site.

Bats of the Savannah River Site
18 Bats of the Savannah River Site and Vicinity

Table 8—Body measurements of the southeastern myotis bat in the

Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 14 86.6 5.24 6.05 77.0 – 95.0

Tail 14 36.9 2.14 5.81 34.0 – 42.0

Foot 14 9.6 1.08 11.22 7.0 – 11.0

Ear 14 13.9 0.53 3.86 13.0 – 15.0

Tragus 4 6.7 1.50 22.22 5.0 – 8.0

Forearm 14 37.1 1.98 5.34 32.0 – 40.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 12 5.7 0.79 13.91 4.3 – 7.0

a Number of individuals measured.

Southeastern myotis

(Myotis austroriparius)

Morphology and Distribution

The southeastern myotis is a small

brown bat with wooly pelage and

a calcar that lacks a keel (fig. 13)

(Jones and Manning 1989). Of the

three subspecies of southeastern

myotis now recognized, only M. a.

austroriparius occurs in South Carolina

(Hoffmeister 1989). The mass of the

southeastern myotis ranges from 5 to

12 g, and total length averages 91.5

mm (Whitaker and Hamilton 1998).

See table 8 for average body

measurements of individuals collected

in the Southeast.

The southeastern myotis’ range

extends from Louisiana and east Texas

up the Mississippi Alluvial Valley into

southern Indiana and Illinois, and east

along the coast of Georgia and the

Carolinas (Jones and Manning 1989).

Four southeastern myotis from South

Carolina are in museum collections,

Figure 13—Southeastern myotis.

and 18 have been live-captured.

Distribution in South Carolina is

limited to the upper and lower Coastal

Plain (fig. 14). Because of its rarity

there, the State has listed the species

as threatened (South Carolina

Department of Natural Resources

2001) (table 9).

Only two southeastern myotis have

been captured on the Site, one male

and one sex undetermined, captured in

1997 and 2000, respectively (table 7).

None was captured during the 1979

surveys. The Site is on the northern

edge of the species’ range, and

although the two documented recordsPhoto courtesy of John MacGregor.
19

Southeastern myotis

(Myotis austroriparius)

Table 9—Protection status of the bats of the Savannah River Sitea

South Carolina

Species State Protection

ranking status Abundance

Eastern pipistrelle Common

Southeastern myotis S1 ST Rare

Little brown myotis S3 SC Uncommon

Small-footed myotisb S1 ST Rare

Northern long-eared myotisb S4 SC Common

Evening bat Common

Rafinesque’s big-eared bat S2 SE Rare

Silver-haired bat Uncommon

Eastern red bat Common

Seminole bat Common

Northern yellow bat SU SC Rare

Hoary bat SU SC Uncommon

Big brown bat Common

Brazilian free-tailed batb Common

S1 = critically imperiled in State; S2 = imperiled in State; S3 = rare or uncommon;

S4 = apparently secure in State; SU = status undetermined; ST = State threatened;

SC = of concern, State; SE = State endangered.

a Only species of concern are assigned rankings and protection status.

b

 Species not captured on the Savannah River Site.

Source: South Carolina Department of Natural Resources (2001).

Bats of the Savannah River Site

Figure 14—Distribution of the southeastern myotis in South Carolina (Menzel

and others 2003). Symbols denote counties for which records exist. Typical

summer distribution is south of the range line.

probably are not extralimital, it appears

to be rare on the Site.

Roosting

Southeastern myotis roost in caves

(Mumford and Whitaker 1982, Rice

1957), mines (Sealander 1979), trees

(Lowery 1974, Sealander 1979), and

buildings (Lowery 1974, Mumford and

Whitaker 1982, Sealander 1979). Davis

and Rippy (1968) banded a colony of

300 individuals roosting in a fertilizer

plant in Georgia. Only Clark and

others (1998) have investigated the

roosting habits of southeastern myotis

in South Carolina. Roosts were found

in cavities of live tupelo gum trees in a

closed-canopy forest on the Francis

Beidler Forest (Clark and others

1998). Based on this information and

the species’ roosting habits in other

areas of its range, we believe it

probably roosts in tree cavities in

bottomland hardwood and swamp

forests on the Site.

Foraging and Home Range

Southeastern myotis typically forage

near water (Barbour and Davis 1969).

In the winter they prey on arthropods

in the orders Coleoptera and

Lepidoptera and in the family Culicidae

(Zinn 1977). During the summer

months they consume primarily

Coleoptera and Lepidoptera (Zinn

1977). Clark and others (1998)

surveyed too few foraging locations to

determine home-range size, but all

foraging points were in swamp forests.

We know little about the size of the

species’ home range or habitat use on

the Site. However, based on the known

home range of other myotids with

similar morphological characteristics,

we surmise the southeastern myotis’

home range is between 100 and 500 ha.

Data collected during the 2001 AnaBat

surveys suggest that foraging activity

on the Site is concentrated in swamp

forest bordering the Savannah River

(fig. 15).

Effect of Habitat Type and

Stand Age on Flight Activity

Southeastern myotis were recorded

at 12.5 percent of the lake and pond

survey points, 22.9 percent of the
20 Bats of the Savannah River Site and Vicinity

0.0

0.5

1.0

1.5

Lake/pond

Carolina bays

Bottomland hardwood

Forest gaps

Grass/brush

Upland hardwood

Pine-hardwood

Loblolly-slash pine

Longleaf pine

Habitat types

Mean number of calls per

20 minute survey

A

B

Figure 15—(A) AnaBat survey locations where southeastern myotis were detected in surveys conducted during summer 2001 and

(B) a comparison of flight activity levels of southeastern myotis among vegetation community types on the Savannah River Site.
21

Photo courtesy of John MacGregor

Carolina bay survey points, 13.6

percent of the bottomland hardwood

survey points, 26.3 percent of the

forest gap survey locations, 5.1 percent

of the grass-brush survey points, 9.1

percent of the upland hardwood points,

7.1 percent of the pine-hardwood

points, 7.1 percent of the loblolly-slash

survey points, and 2.9 percent of the

longleaf survey locations. Overall, we

recorded this species at 10.2 percent

of the survey locations (fig. 15). Its onsite foraging activity was low (table 4).

The mean number of calls recorded

was less than one per 20-minute

survey in all habitat types (fig. 15).

Activity was highest in Carolina bays,

bottomland hardwood forests, and

forest gaps; and the survey points

where southeastern myotis were

detected were concentrated near

swamp forests along the Savannah

River (fig. 15).

Southeastern myotis’ foraging

activity varied among stands of

different ages (fig. 9). Unlike the

eastern pipistrelle’s, most of this

species’ activity was in stands aged 21

to 40 years (table 5).

Reproduction

Southeastern myotis mate in

autumn; fertilization and parturition

occur in spring (Lowery 1974). In

Florida, Rice (1957) found that the

species entered maternity roosts

around mid-March. Most maternity

roosts were in caves and contained

between 2,000 and 90,000 individuals at

average densities of 1,600 bats per m2

.

Young were born from late April to

mid-May. The southeastern myotis

is the only myotid in South Carolina

that typically gives birth to twins

(Barbour and Davis 1969). Although

no maternity colonies have been

studied in South Carolina, parturition

probably occurs in May and early June

on the Savannah River Site.

Evening bat

(Nyticeius humeralis)

Morphology and Distribution

The evening bat is a small brown

bat that occurs throughout the

Southeastern United States (fig. 16).

Of the three recognized subspecies,

only one, N. h. humeralis, occurs

in South Carolina. Weight ranges from

5 to 14 g, and average total length is

92.7 mm (Whitaker and Hamilton

1998). See table 10 for average body

measurements of individuals collected

in the Southeastern United States.

The evening bat’s range extends

north from the Texas-Mexico border

to Nebraska, and east through

Pennsylvania (Watkins 1972). Museum

collections include 172 specimens from

South Carolina; 93 have been livecaptured in the State. The evening bat

occurs in all physiographic provinces

of South Carolina and is common

statewide (fig. 17) (Watkins 1972).

Bats of the Savannah River Site

Figure 16—Evening bat.
22 Bats of the Savannah River Site and Vicinity

Evening bat

(Nycticeius humeralis)

Table 10—Body measurements of the evening bat in the Southeastern

United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 20 86.8 6.28 7.23 77.0 –102.0

Tail 20 33.8 4.07 12.04 26.0 – 39.0

Foot 20 7.9 1.34 16.97 6.0 – 11.0

Ear 19 11.9 2.02 16.96 6.5 – 14.5

Tragus 20 3.9 1.38 35.14 2.5 – 7.0

Forearm 18 36.2 1.81 4.99 33.0 – 39.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 18 8.0 2.04 25.57 4.3 – 12.4

a Number of individuals measured.

Figure 17—Distribution of the evening bat in South Carolina (Menzel and others

2003). Symbols denote counties for which records exist.
23

Evening bats were captured during

the 1979 survey on the Site (Anon.

1980, Cothran and others 1991). In onsite surveys conducted from 1996

to 2001, it was the most commonly

captured species; we captured 27 adult

females, 21 adult males, 13 juvenile

females, and 6 juvenile males (table 7).

Roosting

No studies have quantitatively

examined winter roosting habits of

evening bats in South Carolina, and

little is known about their winter

roosting habits anywhere. There are,

however, anecdotal accounts of

evening bats roosting in attics in

Legareville, Charleston County, SC,

during the winter.

Menzel and others (2000b) found 61

evening bat summer roost trees on the

Site. The most common tree species

was longleaf pine, but conifer snags in

beaver ponds also were common

(Menzel and others 2000b). Evening

bats roosted either in cavities or under

exfoliating bark. Roosts were in areas

with fewer overstory and understory

trees, greater average canopy height,

lower understory richness and

diversity, lower overstory richness,

less-dense canopies, and greater snag

abundance than in random plots

(Menzel and others 2000b). Evening

bat maternity colonies used roosts in

mature longleaf pine in stands where

the height of the overstory, density of

the canopy, and proportion of the basal

area composed of conifers were

greater than in areas surrounding

roost trees used by solitary evening

bats (Menzel and others 2001a).

Summer roosts have been found

elsewhere in Spanish moss (Jennings

1958), under exfoliating bark (Bailey

1933, Barbour and Davis 1969,

Chapman and Chapman 1990), and in

tree cavities (Barbour and Davis 1969,

Harper 1927). There is one record of

this species roosting in caves (Easterla

1965). Maternity roosts have been

found in buildings (Cope and others

1961, Watkins and Shump 1981) and

tree cavities (Menzel and others

2001a). In the upper and lower Coastal

Plain of Georgia, the species often

shares maternity roosts in buildings

with T. brasiliensis.

3

Foraging and Home Range

Evening bats begin foraging in late

twilight (Lowery 1974). At first

emergence, they forage 13 to 25 m

above the ground, but later in the night

they forage lower (Harper 1927,

Lowery 1974). Harper (1927) reported

that in the Okefenokee Swamp area of

Georgia, evening bats foraged in pine

barrens and over fields. Krishon and

others (1997) found that 76 percent of

the bats’ foraging area on Sapelo

Island, GA, was over slash-loblolly pine

habitat. Evening bats feed on insects in

the orders Hemiptera, Hymenoptera,

Coleoptera, and Lepidoptera

(Mumford and Whitaker 1982, Ross

1961). Carter (1998) found that on the

Site they fed primarily on Coleoptera

and Hymenoptera during early and

midsummer. In late summer, the

species fed on a wider variety of insect

taxa, including Hemiptera, Homoptera,

Coleoptera, and Hymenoptera

(in order of decreasing volume)

(Carter 1998).

The home range of one juvenile and

five adult evening bats tracked on the

Site ranged from 38.7 to 761.0 ha and

averaged 285.3 ha (Carter 1998).

Habitat types contained within the

home ranges included pine forests (59

percent) and bottomland hardwoods

(37 percent) (Carter 1998). Habitat

types were used in the same

proportion as they were available.

The 761.0-ha home range of the

juvenile was much larger than that of

the other five. On Sapelo Island, GA,

the species’ home range was within a

15.1-ha slash-loblolly pine stand

(Krishon and others 1997).

Bats of the Savannah River Site

Effect of Habitat Type and

Stand Age on Flight Activity

We recorded evening bats at 31.3

percent of the lake and pond survey

points, 42.9 percent of the Carolina bay

survey points, 31.8 percent of the

bottomland hardwood survey points,

31.6 percent of the forest gap survey

locations, 40.7 percent of the grassbrush survey points, 27.3 percent of

the upland hardwood points, 14.3

percent of the pine-hardwood points,

10.7 percent of the loblolly-slash

survey points, and 21.4 percent of the

longleaf survey locations. Overall, they

were present on 24.9 percent of all

survey locations (fig. 18). Most bat

activity was within grassy areas,

among bottomland hardwoods, and

over Carolina bays (fig. 18). Whereas

eastern pipistrelle activity was highest

over lakes and ponds, evening bat

activity was highest over Carolina

bays. Unlike the southeastern myotis,

evening bat activity was widespread

throughout the Site.

Stand age also influenced evening bat

activity (table 5). As with the eastern

pipistrelle, activity was highest in

clearcuts and young stands (fig. 9).

Moderate activity occurred in stands >

60 years old, but was low in 21- to 60-

year-old stands (fig. 9).

Reproduction

In the Southeastern United States,

evening bats give birth in May and

June. In the Northeast, parturition may

occur in July (Harper 1927, Watkins

1972). Evening bats arrive at nursery

roosts around the second week of April

(Golley 1966) in South Carolina. In

May, Golley (1966) captured several

pregnant females in the State. Most

bats leave their nursery colonies by

late August (Baker 1965). Young bats

become volant about 20 days after

birth (Schmidly 1991). Menzel and

others (2000b) found many evening

bat maternity colonies and captured

many juvenile evening bats on the

Savannah River Site.

3

 Personal communication. 1999. Jim Ozier,

Senior Wildlife Biologist, Georgia Department of

Natural Resources, Wildlife Resources Division

Nongame-Endangered Wildlife Program, 116

Rum Creek Drive, Forsythe, GA 31029-6518.
24 Bats of the Savannah River Site and Vicinity

0.0

0.5

1.0

1.5

2.0

2.5

3.0

3.5

4.0

Lake/pond

Carolina bays

Bottomland hardwood

Forest gaps

Grass/brush

Upland hardwood

Pine-hardwood

Loblolly-slash pine

Longleaf pine

Habitat types

Mean number of calls per

20 minute survey

A

B

Figure 18—(A) AnaBat survey locations where evening bats were detected in surveys conducted during summer 2001 and

(B) a comparison of flight activity levels of evening bats among vegetation community types on the Savannah River Site.
25

Photo courtesy of John MacGregor

Table 11—Body measurements of Rafinesque’s big–eared bat in the

Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 12 96.1 4.54 4.73 90.0 –104.0

Tail 12 46.4 3.11 6.72 43.0 – 53.0

Foot 11 11.8 2.40 20.40 9.0 – 18.0

Ear 11 32.2 2.60 8.08 28.0 – 35.0

Tragus 5 12.7 2.39 18.80 10.0 – 15.0

Forearm 10 43.4 1.33 3.08 41.0 – 46.0

- - - - - - - - - - - - - - - g - - - - - - - - - - - - - - -

Mass 11 8.6 1.05 12.15 7.5 – 11.1

a Number of individuals measured.

Rafinesque’s big-eared bat

(Corynorhinus rafinesquii)

Morphology and Distribution

Rafinesque’s big-eared bat is small,

and its most obvious attribute is its

extremely large ears (fig. 19). Both

subspecies of Rafinesque’s big-eared

bat (C. rafinesquii rafinesquii and

C. r. macrotis) occur in South Carolina

(Jones 1977). Weight ranges from

7 to 10 g, and average total length is

99.5 mm (Whitaker and Hamilton

1998). See table 11 for average body

measurements of individuals collected

in the Southeastern United States.

Rafinesque’s big-eared bat is found

throughout the Southeast. Its range

extends south from southern Virginia,

along the Atlantic Coast into Florida,

and west into Oklahoma and east

Texas (Jones 1977). Museum

collections include 65 specimens

from South Carolina; 30 individuals

have been live-captured in the State.

Its distribution in the South extends

through the upper and lower Coastal

Plain and the Blue Ridge (fig. 20). It is

listed as endangered in South Carolina

(table 9) (South Carolina Department

of Natural Resources 2001).

Two Rafinesque’s big-eared bats

collected on the Site are reposited in

the University of Georgia Museum of

Natural History (Cothran and others

1991). Between 1996 and 2000, we

captured seven adult females and

one adult male (table 7).

Figure 19—Rafinesque’s big-eared bat.

Bats of the Savannah River Site
26 Bats of the Savannah River Site and Vicinity

Rafinesque's big-eared bat

(Corynorhinus rafinesquii)

(Menzel and others 2001b). They

selectively foraged there in saplingstage pine stands (Menzel and others

2001b). The results of this study stand

in stark contrast to the results of a

study conducted by Clark and others

(1998) on the Francis Beidler Forest

in central South Carolina, where the

home range of three individuals was

between 64 and 89 ha, primarily in

swamp forests. On-site, this bat

probably forages in a wide variety of

habitats. However, foraging activity

may be concentrated in mature

bottomland hardwood and swamp

forests, brushy communities, and 3-

to 5-year-old pine plantations.

Fecal pellet analyses in South

Carolina indicated the primary taxa

consumed by Rafinesque’s big-eared

bats was Lepidoptera (Menzel and

others 2002b). Ellis (1993) collected

fecal pellets from the roost of a

maternity colony in North Carolina

and found that the diet was exclusively

composed of Lepidopterans (67

percent) and Dipterans (33 percent).

Most of the flies eaten (94 percent)

were tabanids (family Tabanidae).

Effect of Habitat Type and

Stand Age on Flight Activity

Because of the low intensity of the

echolocation calls produced, activity

patterns of the Rafinesque’s big-eared

bat cannot be monitored reliably with

bat detectors. We did not attempt to

determine the effect of habitat type

or stand age on the flight activity of

this species.

Reproduction

Rafinesque’s big-eared bats seem to

mate in autumn (Hoffmeister and

Goodpaster 1963). Animals with

swollen testes have been found in

August (Hall 1963), and reproductively

active males were captured in areas

around the Site in September (Menzel

and others 2001b). Females enter

maternity roosts in May. Each of two

maternity colonies found after

parturition by Hall (1963) contained

about 50 individuals. The species’

typical litter size is one. Young are

born in late May and early June

(Barbour and Davis 1969, Hall 1963).

Figure 20—Distribution of the Rafinesque’s big-eared bat in South Carolina

(Menzel and others 2003). Symbols denote counties for which records exist.

Roosting

The Rafinesque’s big-eared bat’s

winter roosting habits are not well

known. It has been found roosting in

abandoned buildings throughout the

year (Clark 1990). Winter roosts also

have been found near cave entrances

(Barbour and Davis 1969).

During the summer, Rafinesque’s

big-eared bats roost in abandoned

buildings, hollow trees, and under bark

(Barbour and Davis 1969, Hall 1963,

Jones 1977, Lowery 1974, Menzel and

others 2001b). Typically, summer

roosts are located in the twilight zone

of caves and mines or in dimly lit

sections of buildings (Whitaker and

Hamilton 1998). Menzel and others

(2001b) found male big-eared bats

roosting in abandoned houses in Aiken

County, SC, approximately 8 km

northwest of the Savannah River Site.

We have found some of them roosting

in an abandoned barn on the Site. They

probably roost under bridges, in tree

cavities, and in abandoned structures

near bottomland and swamp forests

on the Site. Clark (1990) described

roosting sites in North Carolina as

“predominately frame, one-story

[abandoned] homes with plaster walls

and large attics.” Lance and others

(2001) found them roosting under

32 girder-type bridges in Louisiana.

Bridges probably are an important

component of the species’ roosting

habitat in South Carolina. Future

surveys should determine which

bridges on the Site serve as roosts for

this species. Lance and others (2001)

found big-eared bats roosting in hollow

blackgum trees. Based on unpublished

museum records, several hundred

were reported roosting in a house on

Belle Isle in Georgetown County, SC.

Clark and others (1998) found some

roosting in tree cavities of water tupelo

in the Francis Beidler Forest in South

Carolina. Most roosts were within

basal cavities. They also found a

1-night roost in a dead bald cypress.

Foraging and Home Range

We know little about the foraging

habits of Rafinesque’s big-eared bats.

They do not begin to forage until after

dark and are agile fliers (Schmidly

1991). In a study conducted in Aiken

County, SC, the home-range size of

male big-eared bats ranged from 23.9

to 260.5 ha and averaged 93.2 ha
27

Photo courtesy of John MacGregor

Bats of the Savannah River Site

In South Carolina, maternity colonies

have been found in abandoned

buildings. Parturition probably

occurs in May and June on the

Savannah River Site.

Silver-haired bat

(Lasionycteris noctivagans)

Morphology and Distribution

The silver-haired bat is a small bat

with silvery-white frosting, and it

occurs throughout southern Canada

and most of the United States (fig. 21).

Weight ranges from 7 to 16 g, and

average total length is 99.7 mm

(Whitaker and Hamilton 1998).

See table 12 for average body

measurements of individuals collected

in the Southeastern United States.

The silver-haired bat reaches the

southern limit of its range in the

Southeast (Kunz 1982). Museum

collections include 26 silver-haired

bats from South Carolina. The species

is distributed statewide (fig. 22),

and individuals have been captured

in all four of South Carolina’s

physiographic provinces. However,

its distribution in the State probably

differs among seasons, and many of

the individuals may have been

captured while migrating.

Although one individual collected on

the Site is reposited in the University

of Georgia Museum of Natural History,

none was captured during the 1979 or

1996 to 2001 surveys. The species is

migratory (Barclay 1984a, Kunz 1982).

Silver-haired bats probably are rare

on the Site between late autumn and

early spring and may not occur there

in the summer.

Roosting

Roosting habits of the silver-haired

bat in South Carolina are unknown.

In other States, they are known to

hibernate in mines (Baker 1965, Layne

1958, Pearson 1962), caves (Baker

1965, Beer 1956, Turner 1974), rock

crevices (Frum 1953), buildings (Clark

1993, Frum 1953), and trees (Cowan

1933, Jackson 1961). Summer roosts

have been found in tree cavities and

under bark (Betts 1996, Mattson and

others 1996, Parsons and others 1986,

Vonhof 1996). Because it is unlikely

that silver-haired bats are on-site

during the summer, the limited

knowledge of the species’ summer

roosting habits in the Southeast is of

little concern.

Foraging and Home Range

Jones and others (1973) and Kunz

(1973) found that silver-haired bats

typically have two peaks of foraging

activity per night—3 and 7 hours after

sunset. Barclay (1984b) found that

they forage throughout the night,

typically in coniferous or mixed

deciduous forests that are near

water (Jones 1965, 1966; Kunz

1973). They fly slowly and erratically

when foraging (Hayward and Davis

1964, van Zyll de Jong 1985). Food

items include Lepidopterans,

Homopterans, Dipterans,

Hemipterans, Hymenopterans,

Coleopterans, and Neuropterans

Figure 21—Silver-haired bat.
28 Bats of the Savannah River Site and Vicinity

Silver-haired bat

(Lasionycterius noctivagans)

Table 12—Body measurements of the silver-haired bat in the

Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Female

Total 3 109.0 1.73 1.59 108.0 –111.0

Tail 3 41.0 4.00 9.76 37.0 – 45.0

Foot 3 8.7 2.08 24.02 7.0 – 11.0

Ear 3 15.0 2.65 17.64 12.0 – 17.0

Tragus 2 5.2 2.47 47.14 3.5 – 7.0

Forearm 3 43.0 1.00 2.33 42.0 – 44.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 3 10.4 0.85 8.20 9.5 – 11.2

- - - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Male

Total 8 95.8 6.84 7.14 81.0 –102.0

Tail 8 39.5 2.20 5.58 35.0 – 42.0

Foot 8 7.31 0.88 12.09 7.0 – 9.5

Ear 8 14.6 1.53 10.46 11.0 – 16.0

Tragus 8 4.0 0.89 22.16 3.0 – 6.0

Forearm 8 40.9 10.9 2.68 43.0 – 40.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 7 8.6 1.58 18.39 5.2 – 9.9

Measures that differed significantly between the sexes (P ≤ 0.05). a Number of individuals measured.

(Black 1974; Whitaker and others

1977, 1981). The species’ foraging

habits in South Carolina are not

known. Because silver-haired bats

probably do not reside on-site during

the summer, the limited knowledge

about its foraging behavior in the

Southeast is of little concern.

Effect of Habitat Type and Stand

Age on Flight Activity

We did not record any calls of silverhaired bats during the 2001 AnaBat

survey. Although they may occur

on-site during the winter months,

it is unlikely that they are summer

residents there.

Reproduction

The reproductive habits of silverhaired bats are not well known.

Similar to other vespertilionid bats,

they probably mate in the autumn

(van Zyll de Jong 1985). Ovulation

occurs in April and May (Druecker

1972). They give birth to one or two

young in June or July (Easterla and

Watkins 1970, Kunz 1971, Merriam

1884). It is unlikely that parturition

occurs on the Savannah River Site.

Figure 22—Distribution of the silver-haired bat in South Carolina (Menzel

and others 2003). Symbols denote counties for which records exist.
29

from 9.5 to 16 g, and its average total

length is 112.3 mm (Whitaker and

Hamilton 1998). See table 13 for

average body measurements of

individuals collected in the Southeast.

The range of the eastern red bat

extends south from southern Canada

into Argentina and Chile (Shump and

Shump 1982a). Museum collections

include 108 specimens from South

Carolina; 145 have been live-captured

in the State. The species is abundant in

all four physiographic provinces of

Eastern red bat

(Lasiurus borealis)

Morphology and Distribution

The eastern red bat is medium

sized and brick red (males) or brick

red with white frosting (females); it

occurs throughout forested regions of

the Eastern United States (fig. 23)

(Shump and Shump 1982a). Of the

three recognized subspecies of eastern

red bats, only one, L. b. borealis, occurs

in South Carolina. Its mass ranges

Photo courtesy of John MacGregor

South Carolina and is common

throughout the State (fig. 24).

One eastern red bat was captured at

Rainbow Bay during the 1979 survey

(Anon. 1980). Between 1996 and 2001,

we captured 30 adult females, 8 adult

males, 11 juvenile females, and 5

juvenile males (table 7). Based on our

capture data, the eastern red bat is the

second most abundant bat species on

the Site.

Roosting

Eastern red bats migrate to the

Southern United States to hibernate.

Generally, winter roosts include tree

branches and leaf clusters (Barbour

and Davis 1969). The species is well

adapted for winter survival outside of

caves and may become active on warm

winter nights (Barbour and Davis

1969). Winter roosting habits in South

Carolina have not been determined. As

with the other species, we know little

of the relative abundance of eastern

red bats on the Savannah River Site

during the winter or of its winter

roosting habits.

In the summer, eastern red bats

most commonly are found roosting on

small branches and leaf petioles in the

crowns of deciduous trees (Barbour

and Davis 1969), but they also have

been found in woodpecker cavities

(Fassler 1975) and caves (Myers

1960). Menzel and others (1998) found

64 eastern red bat roost trees on the

Savannah River Site. Roosts typically

were in hardwood trees larger in

diameter at breast height (d.b.h.) and

taller than surrounding trees. The

roost trees were in areas with greater

overstory tree height and basal area,

denser overstory canopies, and greater

overstory and understory diversity

than random plots (Menzel and others

2000b). Twenty tree species were used

as roosts on the Site, but sweetgum and

red maple were the most common

(Menzel and others 2000b). The

eastern red bat moves to new roost

trees often, spending an average of 1.2

nights at each (Menzel and others

1998). Each individual we tracked

selected roost trees from a small area

of their foraging range (average roost

selection area = 2.6 ha).

Bats of the Savannah River Site

Figure 23—Eastern red bat.
30 Bats of the Savannah River Site and Vicinity

Figure 24—Distribution of the eastern red bat in South Carolina (Menzel and

others 2003). Symbols denote counties for which records exist.

Eastern red bat

(Lasiurus borealis)

Table 13—Body measurements of the eastern red bat in the

Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 36 101.8 7.46 7.33 88.0 –117.0

Tail 36 45.8 4.59 10.01 36.0 – 56.0

Foot 36 7.9 1.36 17.19 6.0 – 10.0

Ear 32 11.3 1.28 11.28 8.0 – 14.0

Tragus 26 4.3 1.29 29.94 3.0 – 8.0

Forearm 33 39.5 2.31 5.85 34.0 – 44.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 27 10.0 1.52 15.15 8.0 – 13.0

a Number of individuals measured.

Foraging and Home Range

Eastern red bats typically begin

foraging 1 to 2 hours after sunset,

although some individuals may

begin earlier and continue foraging

throughout the night (Kunz 1973).

There is some evidence suggesting

they exhibit bimodal foraging activity

patterns (Carter and others 1999a).

The species forages at temperatures

as low as 7 oC (LaVal and LaVal 1979).

In South Carolina, eastern red bats

feed mostly on Coleoptera and

Hemiptera during early summer

(Carter 1998), but in mid- and late

summer they feed on a more diverse

assemblage of taxa, including

Coleoptera, Hemidoptera, Lepidoptera,

Homoptera, and Hymenoptera

(Carter 1998). Elsewhere, remains of

Lepidoptera, Homoptera, Coleoptera,

Hymenoptera, and Diptera have been

found in the stomachs of eastern red

bats (Mumford 1973, Ross 1967,

Whitaker 1972).

The home range of five eastern red

bats tracked on-site ranged from 125.8

to 878.5 ha and averaged 453.2 ha

(Carter 1998). Habitat types within

its home range included bottomland

hardwoods (55 percent), pine stands

(40 percent), and upland hardwoods

(5 percent) (Carter 1998). This species

did not appear to forage selectively in

specific habitat types; they used each

in proportion to its availability.

Effect of Habitat Type and Stand

Age on Flight Activity

Because it is difficult to differentiate

calls of eastern red and Seminole bats,

we included both calls in a single

category. Although data collected in

previous studies suggest eastern red

bats are more likely to forage in

bottomland forests than Seminole bats

(Menzel 1998), pooling the calls made

it impossible to distinguish foraging

habitat use of the two. We recorded

calls at 56.3 percent of the lake and

pond survey points, 71.4 percent of

the Carolina bay survey points, 45.5

percent of the bottomland hardwood

survey points, 68.4 percent of the

forest gap survey locations, 62.7

percent of the grass-brush survey

points, 45.5 percent of the upland
31

Figure 25—(A) AnaBat survey locations where eastern red/Seminole bats were detected in surveys conducted during summer 2001, and

(B) a comparison of flight activity levels of eastern red/Seminole bats among vegetation community types on the Savannah River Site.

0

1

2

3

4

5

6

7

8

9

Lake/pond

Carolina bays

Bottomland hardwood

Forest gaps

Grass/brush

Upland hardwood

Pine-hardwood

Loblolly-slash pine

Longleaf pine

Habitat types

Mean number of calls per

20 minute survey

A

B

Bats of the Savannah River Site
32 Bats of the Savannah River Site and Vicinity

Figure 26—Seminole bat.

Photo courtesy of J. Scott Altenbach

hardwood points, 50.0 percent of the

pine-hardwood points, 36.4 percent of

the loblolly-slash survey points, and

47.1 percent of the longleaf survey

locations. Overall, the species group

was present on 49.7 percent of the

survey locations (fig. 25). The group’s

wide distribution across habitat types

and their high occurrence rate

probably resulted from our inclusion

of both species. The effect of pooling

would have an even greater effect on

these metrics if they commonly

foraged in different habitats. If

Seminole bats usually roost and forage

in upland habitats and eastern red bats

in bottomland forests, such pooling

would result in a group common to

both upland and bottomland

communities. Also, because the

eastern red and Seminole bats are

the second and third most common

species on the Site, respectively,

it is not surprising that this group

was detected on half of all 449

survey locations.

In addition to being commonly

detected, the group’s activity levels

were relatively high in all habitat types

(fig. 25). Most activity was detected

over Carolina bays and grass-brush

habitat (table 4).

Eastern red and Seminole bat

activity also seemed to be less affected

by stand age than that of other species

(fig. 9). Although activity was greatest

within clearcuts, it was relatively

constant among stands aged 4 years or

older (table 5).

Reproduction

Eastern red bats undergo delayed

fertilization. Mating occurs in August

and September, and fertilization in

spring (Glass 1966, Whitaker and

Hamilton 1998). Copulation sometimes

occurs in flight (Stuewer 1948).

Estimated gestation is between 80 and

90 days (Jackson 1961). Its young are

born in late May and June (Lowery

1974), and lactation lasts about 38 days

(Kunz 1971). The average embryo

count is 3.2 (Shump and Shump

1982a). Members of the genus

Lasiurus are the only bats that

regularly give birth to more than two

progeny (Nowak 1994). Although no

one has studied the reproductive

habits of eastern red bats in South

Carolina, we have captured juveniles

on the Site, and it is likely that they

commonly raise their young on-site.

Seminole bat

(Lasiurus seminolus)

Morphology and Distribution

The Seminole bat is a medium sized,

mahogany colored bat common

throughout the Deep South (fig. 26).

The Seminole bat is monotypic. Its

mass ranges from 9 to 14 g, and

its average total length is 111 mm

(Whitaker and Hamilton 1998).

See table 14 for average body

measurements of individuals collected

in the Southeast.

Although a few “wandering”

individuals have been captured in New

York (Layne 1955) and Pennsylvania

(Poole 1949), this species typically is

found south from the southeastern tip

of Virginia to Florida, and west along
33

Figure 27—Distribution of the Seminole bat in South Carolina (Menzel and others

2003). Symbols denote counties for which records exist. Typical summer

distribution is south of the range line.

Seminole bat

(Lasiurus seminolus)

Table 14—Body measurements of the Seminole bat in the

Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Female

Total 16 106.1 6.65 6.27 89.0 –116.0

Tail 16 43.7 3.91 8.95 38.0 – 53.0

Foot 16 8.6 0.81 9.51 6.0 – 9.0

Ear 15 12.2 0.98 8.01 11.0 – 14.0

Tragus 14 6.5 1.57 24.09 4.0 – 9.0

Forearm 16 42.3 0.94 2.23 41.0 – 45.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 14 12.7 1.80 14.15 8.5 – 15.0

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Male

Total 18 101.8 5.59 5.49 93.0 –115.0

Tail 18 43.9 3.89 8.85 38.0 – 52.0

Foot 17 8.1 1.25 15.49 5.0 – 9.0

Ear 16 12.0 1.15 9.62 10.0 – 14.0

Tragus 17 6.1 1.49 24.44 3.0 – 8.0

Forearm 18 40.5 1.40 3.45 38.0 – 43.5

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 12 9.8 1.18 12.09 7.8 – 11.0

Measures that differed significantly between the sexes (P ≤ 0.05). a Number of individuals measured.

Roosting

In the winter, Seminole bat roosts

commonly are found in oak hammock

communities in Spanish moss

(Constantine 1958), but the species’

winter roosting habits in South

Carolina are not known.

Seminole bats also commonly roost

in Spanish moss during the summer

months (Barbour and Davis 1969).

Although one summer roost was

located under loose bark, this species

typically does not use bark roosts

(Sealander 1979). On the Savannah

River Site, summer roosts usually are

in the terminal branches of pine limbs

in communities dominated by pines

(Menzel and others 1998). Although 10

tree species were used as roosts, most

(86 percent) were pines, primarily

loblolly (Menzel and others 2000b).

Roosts tended to be in taller, larger

diameter trees than the surrounding

overstory (Menzel and others 2000b).

Roosts were found in areas with higher

basal area, lower understory species

richness, and less Spanish moss than

random plots (Menzel and others

2000b). On-site, Seminole bats move

Bats of the Savannah River Site

the Gulf Coast into east Texas (Wilkins

1987). Museum collections include 42

Seminole bats from South Carolina; 58

have been live-captured in the State.

Distribution of the Seminole bat in

South Carolina includes the upper and

lower Coastal Plain (fig. 27). They have

been captured in the Piedmont and

Blue Ridge, but their distribution in

surrounding States suggests they

rarely occur above the fall line.

The Seminole bat was one of the

most commonly captured bat species

during the 1979 survey (Anon. 1980).

In surveys conducted between 1996

and 2001, we captured six adult

females, seven adult males, six juvenile

females, and three juvenile males

(table 7). Based on surveys conducted

on the Site, the Seminole bat is the

third most common species, exceeded

in abundance only by evening and

eastern red bats.
34 Bats of the Savannah River Site and Vicinity

Photo courtesy of John MacGregor

Figure 28—Hoary bat.

to new roost trees often, spending

an average of 1.7 days in each. On

average, they select new roost trees

within about a 0.25-ha portion of their

home range.

Foraging and Home Range

Although Seminole bats commonly

forage at treetop level (Barbour and

Davis 1969), two reports document

the species gleaning prey from

leaf surfaces (Barbour and Davis

1969, Sherman 1935). Sherman

(1939) found the remains of

Homopterans (Jassidae), Dipterans

(Dolichopodidae, Muscidae), and

Coleopterans (Scolytidae) in a single

stomach. Zinn (1977) reported

that Coleoptera, Odonata, and

Hymenoptera were important

food items. Carter (1998) found

that Seminole bats fed primarily

on Hymenoptera, Coleoptera, and

Lepidoptera on the Site.

Radio tracking on the Site indicated

the home-range size of five Seminole

bats to be between 189.2 and 704.4 ha,

with an average of 423.8 ha (Carter

1998). Habitat types in the home-range

areas included pine forests (55

percent), bottomland hardwoods

(35 percent), and upland hardwoods

(11 percent) (Carter 1998).

Effect of Habitat Type and

Stand Age on Flight Activity

Because it is hard to differentiate

calls of eastern red and Seminole bats,

we included both calls in a single

category. So far, using acoustical

monitoring to determine the effect of

habitat type and stand age on the flight

activity of this species has not been

possible. See the eastern red bat

species account for a description of the

effect of habitat type and stand age on

the flight activity of the eastern red/

Seminole bat group.

Reproduction

Pregnant Seminole bats have been

collected between early May and midJune (Barkalow 1948, Coleman 1950,

Jennings 1958, Moore 1949).

Parturition usually occurs in June and

July, but the species’ reproductive

habits in South Carolina are not

known. However, we captured juvenile

Seminole bats on the Savannah River

Site and tracked adult females to

roosts occupied by their young.

Seminole bats raise their progeny

on the Site.

Hoary bat

(Lasiurus cinereus)

Morphology and Distribution

The hoary bat is the largest bat

species found in South Carolina. It has

frosted pelage and cream-colored ears

with a distinct back rim (fig. 28).

Of three subspecies of the hoary bat,

one, L. c. cinereus, occurs in the State

(Shump and Shump 1982b). Its mass

ranges from 18 to 38 g, and its average

total length is 134.8 mm (Whitaker

and Hamilton 1998). See table 15 for

average body measurements of

individuals collected in the Southeast.
35

Figure 29—Distribution of the hoary bat in South Carolina (Menzel and others

2003). Symbols denote counties for which records exist.

Hoary bat

(Lasiurus cinereus)

Table 15—Body measurements of the hoary bat in the Southeastern

United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 7 132.5 12.72 9.60 106.0 – 146.0

Tail 7 58.0 4.73 8.15 51.0 – 63.0

Foot 6 11.5 1.52 13.19 10.0 – 14.0

Ear 7 14.9 2.79 18.81 10.0 – 18.0

Tragus 5 7.3 1.30 17.86 5.5 – 9.0

Forearm 6 55.2 1.94 3.52 53.0 – 58.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 6 21.2 2.40 11.32 19.0 – 24.9

a Number of individuals measured.

The hoary bat has a more extensive

distribution than any other bat species

in South Carolina. Its range extends

south from northern Canada near the

tree line through Central America. The

southern limit of its range extends into

Argentina and Chile (Shump and

Shump 1982b). Museum collections

include two hoary bats from South

Carolina; three have been live-captured

in the State. The species’ distribution

in South Carolina (fig. 29) includes all

four physiographic provinces, and its

status is listed as undetermined (table

9) (South Carolina Department of

Natural Resources 2001). Because of

its migratory patterns, it probably is

rare in the State during summer.

We captured three hoary bats (two

females and one sex undetermined) on

the Site from 1996 to 2001 (table 7).

Whitaker and Hamilton’s (1998)

summer range map for hoary bats

does not include South Carolina.

Although our surveys indicate that

they do occur on the Site in summer,

they are not abundant.

Roosting

Although the hoary bat’s winter

roosts typically are found in tree

foliage, they also have been found in

tree cavities and squirrel nests

(Constantine 1966, Cowan and Guiguet

1965, Neill 1952). The species typically

does not roost in caves (Mumford

1953, Myers 1960).

Little is known about the hoary bat’s

summer or winter roosting habits in

South Carolina, and nothing is known

about its roosting requirements on

the Site.

Foraging and Home Range

Hoary bats begin foraging in late

evening (Barbour and Davis 1969),

sometimes establishing foraging

territories (Barclay 1984a). Prey items

include Coleopterans, Dipterans,

Orthopterans, Isopterans, Odonatans,

Hymenopterans, and Lepidopterans

(Black 1972, 1974; Ross 1967;

Whitaker 1972; Whitaker and others

1977), but it seems to prefer

Lepidopterans (Black 1972, Ross

1967). Its foraging habits in South

Bats of the Savannah River Site
36 Bats of the Savannah River Site and Vicinity

Figure 30—(A) AnaBat survey locations where hoary bats were detected in surveys conducted during summer 2001, and

(B) a comparison of flight activity levels of hoary bats among vegetation community types on the Savannah River Site.

0.0

0.5

1.0

1.5

2.0

2.5

3.0

Lake/pond

Carolina bays

Bottomland hardwood

Forest gaps

Grass/brush

Upland hardwood

Pine-hardwood

Loblolly-slash pine

Longleaf pine

Habitat type

Mean number of calls per

20 minute survey

A

B
37

Figure 31—Big brown bat.

Photo courtesy of John MacGregor

Carolina are poorly understood.

However, its diet on the Savannah

River Site probably is similar to its diet

elsewhere in its range.

We have found no information about

the size or spatial characteristics of the

species’ home range.

Effect of Habitat Type and

Stand Age on Flight Activity

We recorded hoary bats on 18.8

percent of the lake and pond survey

points, 5.7 percent of the Carolina bay

survey points, 7.6 percent of the

bottomland hardwood survey points,

2.6 percent of the forest gap survey

locations, 6.8 percent of the grassbrush survey points, none of the

upland hardwood points, none of the

pine-hardwood points, 5.0 percent of

the loblolly-slash survey points, and

1.4 percent of the longleaf survey

locations. Overall, we recorded it on

5.2 percent of all survey locations

(fig. 30). Its activity level was

low throughout the Site, primarily

concentrated in bottomland hardwood

stands (table 4).

Stand age affected hoary bat activity

differently than it affected that of other

bat species on the Site (fig. 9). Based

on the low call frequency, high

wing loading, and moderately high

wing-aspect ratio of the species, we

expected activity would be highest in

clearcuts; yet we recorded no hoary

bat calls there and found most activity

concentrated in stands aged > 60

years (table 5).

Reproduction

Hoary bats probably mate during

autumn migration (Shump and Shump

1982b). Parturition occurs from May

through July (Kurta 1982, McClure

1942, Mumford 1969, Provost and

Kirkpatrick 1952, Whitaker and

Mumford 1972). Because the species

migrates north in spring, parturition

may not occur in South Carolina.

Big brown bat

(Eptesicus fuscus)

Morphology and Distribution

The big brown bat is a medium

sized, nondescript brown bat and is

the third largest in South Carolina (fig.

31). One of the 11 subspecies of big

brown bat, E. f. fuscus, occurs in the

State (Kurta and Baker 1990). Its mass

ranges from 13 to 25 g, and its average

total length is 114.3 mm (Whitaker and

Hamilton 1998). See table 16 for

average body measurements of

individuals collected in the Southeast.

The range of the big brown bat

extends from the Atlantic to the Pacific

Coasts, and south from Canada

through Central America and into

Brazil (Kurta and Baker 1990). It is

found throughout the continental

United States and is the most common

bat species throughout much of its

range. It is most abundant in the

Bats of the Savannah River Site
38 Bats of the Savannah River Site and Vicinity

Figure 32—Distribution of the big brown bat in South Carolina (Menzel and

others 2003). Symbols denote counties for which records exist.

Big brown bat

(Eptesicus fuscus)

Table 16—Body measurements of the big brown bat in the

Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Female

Total 23 114.1 6.38 5.59 100.0 – 122.0

Tail 23 44.2 3.44 7.78 34.0 – 49.0

Foot 23 9.4 1.55 16.46 6.5 – 13.0

Ear 23 17.4 1.99 11.49 12.0 – 20.0

Tragus 20 5.7 1.43 25.21 3.0 – 9.0

Forearm 23 45.5 9.44 20.78 4.0 – 52.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 21 17.2 4.04 23.54 11.0 – 27.0

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Male

Total 16 107.6 6.59 6.13 97.0 – 119.0

Tail 16 41.7 4.67 11.18 30.0 – 49.0

Foot 16 9.7 1.08 11.13 8.0 – 12.0

Ear 15 16.4 1.80 11.00 13.0 – 19.0

Tragus 14 6.4 1.11 17.40 5.0 – 8.0

Forearm 15 46.6 2.53 5.43 40.0 – 50.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 16 12.8 3.12 24.33 7.0 – 18.8

Measures that differed significantly between the sexes (P ≤ 0.05). a Number of individuals measured.
39

deciduous forest biome (Kurta and

others 1989). Museum collections

include 15 big brown bats from South

Carolina; 18 have been live-captured

there. It is common throughout the

State (fig. 32) and is found in all four

physiographic provinces.

Despite its apparent abundance in

South Carolina, this species was not

documented in the 1979 survey, and

between 1996 and 2001 we captured

only six (two adult females, three

adult males, and one juvenile female)

(table 7). The relatively few captures

on the Site probably was due to the

scarcity of preferred roosts. Although

it uses many different structures

(see Roosting section), summer

roosts are most often found in the

attics of houses.

Roosting

Big brown bats are one of the most

studied in North America. Their

hibernacula have been found in

mines, caves, rock crevices, storm

sewers, and buildings (Barbour

and Davis 1969, Beer 1955, Fenton

1972, Mills and others 1975, Mumford

1958, Nagorsen 1980, Phillips 1966,

Quay 1949).

Summer roosts usually are found in

hollow oak and American beech

(Fagus grandifolia Ehrh.), but most

commonly the species uses humanmade structures such as barns or

houses (Barbour and Davis 1969,

Christian 1956, Kurta 1982). In early

August 1996, Menzel (1998) and

Carter (1998) radio tracked two big

brown bats in a bottomland hardwood

swamp to a maternity roost in a hollow

bald cypress snag along the Savannah

River. Other than this anecdotal

account, no studies have investigated

the species’ summer or winter roosting

habits in South Carolina.

Foraging and Home Range

Big brown bats typically begin

foraging within the first hour after

sunset (Kurta 1982, Phillips 1966).

They feed on Isoptera, Hemiptera,

Homoptera, Hymenoptera,

Lepidoptera, and Diptera (Freeman

1981, Menzel and others 2000a, Ross

1967), although most of their diet is

composed of beetles (Hamilton 1933;

Menzel and others 2000a; Phillips

1966; Whitaker 1972, 1995). Menzel

and others (2000a) reported that

during the reproductive period,

females selectively forage on

Coleopterans and eat fewer

Lepidopterans, Dipterans, and

Hymenopterans, based on the

availability of such insect orders

in the foraging area. The species’

foraging habits in South Carolina

are not known.

To date, little quantitative

information has been gathered about

this species’ foraging habitat selection.

Furlonger and others (1987) found that

they were generalists when selecting

habitat. There are many reports of

foraging around urban streetlights,

e.g., Geggie and Fenton 1985. In the

Georgia Piedmont, Menzel and others

(2001c) found that big brown bats

selectively foraged in rural rather than

urban areas, and less often in

agricultural fields and clearcuts than in

hardwood and pine forests. Menzel

and others (2001c) reported that the

average home-range size was 2906 ha;

Brigham (1991) reported their

maximum foraging distance to be 4.4

km. Both these sets of data suggest

large foraging ranges. Information

about the foraging behavior of this

species in other regions of its range

suggests that foraging activity on the

Site would not be concentrated in a

single habitat type, but would be

dispersed across a variety of types.

The species probably forages in

both upland and bottomland habitats

on the Savannah River Site.

Effect of Habitat Type and

Stand Age on Flight Activity

Big brown bats were recorded at

25.0 percent of the lake and pond

survey points, 20.0 percent of the

Carolina bay survey points, 15.2

percent of the bottomland hardwood

survey points, 5.3 percent of the forest

gap survey locations, 18.6 percent of

the grass-brush survey points, none

of the upland hardwood points, 7.1

percent of the pine-hardwood points,

7.9 percent of the loblolly-slash survey

points, and 10.0 percent of the longleaf

survey locations. We recorded them on

11.8 percent of all survey locations (fig.

33). Activity was concentrated over

lakes and ponds, grass-brush, and in

bottomland hardwoods (table 4).

Foraging activity appeared to be

unaffected by stand age (fig. 9).

Reproduction

Mating occurs in September and

March (Mumford 1958, Phillips

1966), and fertilization in spring

(Wimsatt 1944). The species’

reproductive habits are unknown in

South Carolina, but we captured a

juvenile on the Site and tracked two

females to a large (> 30 individuals)

maternity colony located in a bald

cypress snag where Pen Branch flows

into a swamp forest adjacent to the

Savannah River. The species probably

raises its young on-site.

Bats of the Savannah River Site
40 Bats of the Savannah River Site and Vicinity

Figure 33—(A) AnaBat survey locations where big brown bats were detected in surveys conducted during summer 2001, and

(B) a comparison of the flight-activity levels of big brown bats among vegetation community types on the Savannah River Site.

0.0

0.5

1.0

1.5

Lake/pond

Carolina bays

Bottomland hardwood

Forest gaps

Grass/brush

Upland hardwood

Pine-hardwood

Loblolly-slash pine

Longleaf pine

Habitat types

Mean number of calls per

20 minute survey

A

B
41

Photo courtesy of John MacGregor

Figure 34—Little brown bat.

Table 17—Body measurements of the little brown bat in the

Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 10 87.4 4.96 5.67 80.0 – 95.0

Tail 10 37.5 3.03 8.08 32.0 – 41.0

Foot 9 8.1 0.73 9.02 6.5 – 9.0

Ear 10 13.5 1.08 8.00 12.0 – 15.0

Tragus 9 5.7 1.30 22.92 4.5 – 8.0

Forearm 7 37.0 2.02 5.46 34.0 – 40.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 10 6.5 1.09 16.62 5.5 – 9.3

a Number of individuals measured.

Bats of South

Carolina in Areas

Neighboring the

Savannah River Site

FAMILY VESPERTILIONIDAE

Little brown bat

(Myotis lucifugus)

Morphology and Distribution

T he little brown bat is a small

brown bat with glossy pelage

that occurs in some areas of

South Carolina (fig. 34). Of the six

subspecies currently recognized, only

M. l. lucifugus occurs in the State

(Fenton and Barclay 1980). Its mass

ranges from 4 to 8 g, and its average

total length is 89.2 mm (Whitaker and

Hamilton 1998). See table 17 for

average body measurements of

individuals collected in the Southeast.

The little brown bat is common

throughout much of the United States.

In the Eastern States, the southern

limit of its range reaches into northern

portions of South Carolina, Georgia,

Alabama, and Mississippi (Fenton

and Barclay 1980). Museum

collections include three in South

Carolina; three have been live-captured

there. Although its typical distribution

in the State is limited to the Blue

Ridge (fig. 35), captures have been

confirmed in Beaufort County on

the lower Coastal Plain (Davis and

Rippy 1968). South Carolina has listed

it as uncommon or rare (South

Bats of South Carolina in Areas Neighboring the Savannah River Site
42 Bats of the Savannah River Site and Vicinity

Figure 35—Distribution of the little brown bat in South Carolina (Menzel and

others 2003). Symbols denote counties for which records exist. Records south of

the range line likely are extralimital, because the primary distribution of this

species is in the Blue Ridge and northward.

Table 18—Body measurements of the eastern small-footed myotis in

the Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 2 78.0 2.83 3.63 76.0 – 80.0

Tail 2 33.0 1.41 4.28 32.0 – 34.0

Foot 2 6.0 1.41 23.57 5.0 – 7.0

Ear 2 14.0 0. 0. 14.0 – 14.0

Tragus 2 5.0 NA NA 5.0 – 5.0

Forearm 1 31.0 1.41 4.56 30.0 – 32.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 2 4.2 0.07 1.66 4.2 – 4.3

a Number of individuals measured.

Carolina Department of Natural

Resources 2001).

One female little brown bat was

captured on the Site in 1996 (table 7).

Because the Site is south of the species’

typical range, that single record

probably is extralimital. The species

probably does not occur

there regularly.

Roosting

Little is known about the roosting

habits of little brown bats in South

Carolina. Its hibernacula most

commonly are found in caves and

mines (Baker 1965, Humphrey and

Cope 1976), but no hibernacula have

been found in the State. In summer,

little brown bats roost under rocks, in

piles of wood, and in trees (Fenton and

Barclay 1980). Maternity roosts

commonly are found in buildings

(Davis and Hitchcock 1965, Griffin

1940, Youngman 1975). The summer

or winter roosting habits of this

species in South Carolina have not

been reported.

Foraging and Home Range

Little brown bats emerge shortly

after dusk to feed (Nagorsen and

Brigham 1990). Numerous foraging

habit studies have been conducted

(Anthony and Kunz 1977; Belwood and

Fenton 1976; Buchler 1976; Ross 1961,

1967; Whitaker 1972), but the species’

diet in South Carolina is not known. It

typically forages along riparian areas

(Fenton and Barclay 1980). Griffin and

others (1960) found that when feeding

in an insect swarm, little brown bats

could capture up to 12 fruit flies per

minute. Although the species’ diet is

variable, aquatic insects constitute

most of its forage (Fenton and Barclay

1980). Little is known about the size of

its home range, and nothing is known

about its home range or habitat use in

South Carolina. Because the species

does not occur there regularly, its

foraging behavior should be of little

concern to land managers on the Site.

Effect of Habitat Type and

Stand Age on Flight Activity

We did not record any calls of

little brown bats during the 2001

AnaBat survey.

Reproduction

Little brown bats mate in autumn,

and fertilization occurs in spring when

females leave the hibernacula.

Duration of hibernation typically

decreases with decreasing latitude,

resulting in earlier ovulation and

parturition dates at more southerly

latitudes (Fenton and Barclay 1980).

Parturition occurs in late May and

June (Barbour and Davis 1969,

Hayward 1963, Hoffmeister 1989)

on the Savannah River Site, and it is

unlikely that little brown bats raise

their young there.
43

Photo courtesy of John MacGregor

Figure 36—Eastern small-footed myotis.

Figure 37—Distribution of the eastern small-footed myotis in South Carolina

(Menzel and others 2003). Symbols denote counties for which records exist.

Typical summer distribution is north of the range line.

Eastern small-footed myotis

(Myotis leibii)

Morphology and Distribution

The eastern small-footed myotis

is a small brown bat with long, glossy

pelage and a black mask (fig. 36).

It is the smallest bat found in South

Carolina. This species is monotypic

(van Zyll de Jong 1985). Its mass

ranges from 4 to 6 g, and its average

total length is 77.5 mm (Whitaker

and Hamilton 1998). See table 18

for average body measurements of

individuals collected in the Southeast.

The species’ range is restricted to

northeastern North America, from

extreme northwestern South Carolina

west into Oklahoma and Missouri and

north into the St. Lawrence forest

region of Ontario and Quebec (van Zyll

de Jong 1985). Museum collections

include 42 eastern small-footed myotis

from South Carolina. Within the State,

its range is limited to the Blue Ridge

in the extreme north (fig. 37). The

species appears to be uncommon

Bats of South Carolina in Areas Neighboring the Savannah River Site
44 Bats of the Savannah River Site and Vicinity

Photo courtesy of John MacGregor

Figure 38—Northern long-eared myotis.

throughout its range, and it is listed as

threatened in South Carolina (table 9)

(South Carolina Department of Natural

Resources 2001).

No eastern small-footed myotis’ have

been captured on the Site. Based on

its distribution and roosting-habitat

preferences, i.e., rock outcrops, it is

highly unlikely that this species would

occur on the Site.

Roosting

Roosting habits of eastern smallfooted myotis’ in South Carolina are

not well known. In other States,

hibernacula have been found in caves

and mines (Davis 1955; Fenton 1972;

Hitchcock 1945, 1949; Mohr 1936).

Summer roosts have been found

under rocks and in buildings (Barbour

and Davis 1969, Hitchcock 1955,

Tuttle 1964).

Foraging and Home Range

Eastern small-footed myotis’ begin

foraging shortly after sunset. Average

foraging height is 1 to 3 m above the

ground (van Zyll de Jong 1985), but

little is known about its prey.

Because it is so small, this species is

difficult to track using radiotelemetry.

To lessen the transmitter’s effect on

foraging behavior, transmitter mass

was < 0.4 g. Because of this difficulty,

and because of the species’ rarity,

characteristics of its home range have

not been documented.

Effect of Habitat Type and

Stand Age on Flight Activity

We did not record any eastern smallfooted myotis calls during the 2001

AnaBat survey. This species has not

been captured on the Site, and it is

unlikely to occur there.

Reproduction

Information about eastern smallfooted myotis reproduction is limited to

one record of a maternity colony located

behind the door of a barn in Ontario,

Canada, in summer (Hitchcock 1955).

No maternity colonies have been

found in South Carolina.

Northern long-eared myotis

(Myotis septentrionalis)

Morphology and Distribution

The northern long-eared myotis is

a medium-sized bat with long, glossy

brown pelage (fig. 38) (Fitch and

Shump 1979). Its mass ranges from

5 to 10 g, and its average total length

is 84.1 mm (Whitaker and Hamilton

1998). See table 19 for average body

measurements of individuals collected

in the Southeast.

The northern long-eared myotis

occurs throughout much of the eastern

half of the United States. Its range

extends east from Saskatchewan

to Quebec, Canada, and south to

Florida (Fitch and Shump 1979).

Although there are no museum

specimens from South Carolina, 22

individuals have been live-captured in

surveys. Its distribution in the State is
45

Figure 39—Distribution of the northern long-eared myotis in South Carolina

(Menzel and others 2003). Symbols denote counties for which records exist.

Typical summer distribution is north of the range line.

4 Menzel, M.A. Unpublished data. On file with:

West Virginia University, Division of Forestry,

Percival Hall, Morgantown, WV 26506.

Table 19—Body measurements of the northern long-eared myotis in

the Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 15 85.3 5.09 5.97 75.0 – 92.0

Tail 15 37.6 3.42 9.09 30.0 – 42.0

Foot 15 7.2 0.86 11.97 6.0 – 9.0

Ear 14 15.3 1.88 12.25 11.0 – 17.0

Tragus 12 6.8 1.01 14.74 5.0 – 8.5

Forearm 15 35.1 0.89 2.54 33.5 – 36.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 12 8.0 11.65 144.92 3.6 – 45.0

a Number of individuals measured.

restricted to the Blue Ridge, where the

species is common (fig. 39) (South

Carolina Department of Natural

Resources 2001).

No northern long-eared myotis

have been captured on the Savannah

River Site. Its typical range does not

include the Site, and it is unlikely

any occur there.

Roosting

Hibernacula used by northern longeared myotis have been found in caves

(Griffin 1940, Swanson and Evans

1936), mines (Stones and Fritz 1969),

and a storm sewer (Goehring 1954).

During summer, the species roosts

in buildings (Doutt and others 1966,

Turner 1974), behind shutters

(Mumford 1969), and under the bark

and in the cavities of trees (Menzel

and others 2002d, Mumford and

Cope 1964). Its summer or winter

roosting habits in South Carolina are

not known.

Foraging and Home Range

Northern long-eared myotis forage

shortly after dusk and before dawn

(Barbour and Davis 1969). They forage

around ponds and along hillsides, both

above (Cowan and Guiguet 1965) and

beneath (LaVal and others 1977) the

tree canopy. Little is known about their

diet. Their foraging habits in South

Carolina are not known.

Although this species’ home-range

characteristics in the Southeast

have not been studied, data are

available for the central Appalachians.

In the Allegheny Mountains of West

Virginia, its home ranges are

unimodal, from 43 to 578 ha (mean

216 ha), and include clearcuts,

deferment harvests, streams, road

corridors, and mature second-growth

eastern deciduous forests.4

Effect of Stand Age and

Habitat Type on Flight Activity

We did not record any calls of

northern long-eared myotis during

the 2001 AnaBat survey. We have not

captured any of this species on the

Site, and it is unlikely to occur there.

Reproduction

Northern long-eared myotis copulate

in autumn, and sperm is stored in

the uterus throughout hibernation.

Fertilization occurs in spring (Barbour

and Davis 1969). Parturition occurs in

June, and the young typically become

volant in late July (Easterla 1968, Kunz

1971). Parturition times and maternity

colony habits in South Carolina are

not known.

Bats of South Carolina in Areas Neighboring the Savannah River Site
46 Bats of the Savannah River Site and Vicinity

Figure 40—Northern yellow bat.

Photo courtesy of J. Scott Altenbach

5

 Menzel, M.A.; Krishon, D.M.; Carter, T.C. 1995.

Roosting, foraging, and habitat use by bats of

Sapelo Island, Georgia. Unpublished Tech. Rep.

60 p. On file with: Georgia Department of Natural

Resources, Attn: J. Ozier., Social Circle, GA.

Northern yellow bat

(Lasiurus intermedius)

Morphology and Distribution

The northern yellow bat is a large

yellow bat and is the second largest bat

in South Carolina (fig. 40). Although

two subspecies of it are recognized,

the validity of this designation is a

matter of debate (Whitaker and

Hamilton 1998). Of the two subspecies,

L. intermedius floridanus occurs in

South Carolina. Its mass ranges from

14 to 20 g, and average total length is

126.8 mm (Whitaker and Hamilton

1998). See table 20 for average body

measurements of individuals collected

in the Southeast.

Although one northern yellow bat

was collected in New Jersey (Koopman

1965), typically the species is restricted

to coastal areas of the Southeastern

United States and Central America

(Webster and others 1980). Museum

collections include 11 northern yellow

bats from South Carolina; one has

been live-captured in the State. It

occurs in the lower Coastal Plain, and

its range extends into the upper

Coastal Plain along the Savannah River

(fig. 41). The State has listed the status

of this species as undetermined (table

9) (South Carolina Department of

Natural Resources 2001).

One northern yellow bat was

collected along Upper Three Runs

Creek in 1978. However, none was

captured during the 1979 survey

(Anon. 1980) or the 1996 to 2001

survey. Based on the species’

distribution, it probably is not

a common resident on the

Savannah River Site. The individual

collected along Upper Three Runs

Creek probably represents an

extralimital record.

Roosting

Little is known about the northern

yellow bat’s winter roosting habits in

South Carolina or elsewhere.

Summer roosts have been located

in palm groves (Davis 1960), on

hardwood stems (Rageot 1955), in

pine-oak woodlands (Carter and Jones

1978, Carter and others 1966, Jones

1964, Sherman 1944), and in Spanish

moss (Jennings 1958).5

 Northern

yellow bats usually roost alone;

however, Baker and Dickerman (1956)

reported seeing about 45 individuals

fly from a communal roost under corn

stalks hanging from the side of an old

tobacco curing shed. The species

is not a regular summer resident on

the Site and should be of little

management concern.

Foraging and Home Range

Northern yellow bats typically begin

foraging well before dark (Lowery

1974). They forage 4 to 6 m above the

ground (Barbour and Davis 1969) and

prefer open foraging areas such as

airports, golf courses, and fields

(Jennings 1958). Lowery (1974)

observed them foraging over the

Mississippi River. After the young

become volant, they form feeding
47

Figure 41—Distribution of the northern yellow bat in South Carolina (Menzel and

others 2003). Symbols denote counties for which records exist. Typical summer

distribution is north of the range line.

Table 20—Body measurements of the northern yellow bat in the

Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 5 127.6 3.36 2.63 124.0 – 132.0

Tail 5 52.6 2.51 4.77 49.0 – 55.0

Foot 5 10.4 1.52 14.58 8.0 – 12.0

Ear 3 17.7 2.52 14.24 15.0 – 20.0

Tragus 3 7.7 3.05 39.85 5.0 – 11.0

Forearm 3 51.7 1.15 2.23 51.0 – 53.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 3 20.0 2.29 11.46 17.5 – 22.0

a Number of individuals measured.

aggregations that may consist of

> 100 individuals (Jennings 1958). Such

aggregations primarily are composed

of females. Northern yellow bats

emerge and feed on warm nights

throughout winter (Jennings 1958).

Northern yellow bats feed on

Homoptera, Zygotera, Diptera,

Coleoptera, and Hymenoptera

(Sherman 1939), but their foraging

habits in South Carolina are

not known.

The home range of a northern

yellow bat tracked on Sapelo Island,

GA, was 10.5 ha (Krishon and others

1997). It was located in oak (73

percent) and slash-loblolly pine (25

percent) communities. The average

distance from the roost area center to a

foraging location was 109 m. This is a

relatively small home range compared

to the average home range of other,

smaller lasiurines, e.g., eastern red bat

= 453.2 ha, Seminole bat = 428.8 ha.

This estimate is based on one

individual and may not represent the

species’ average home-range size. No

home range or habitat use studies of

northern yellow bats have been

conducted in South Carolina.

Effect of Habitat Type and

Stand Age on Flight Activity

We did not record any calls of

northern yellow bats during the 2001

AnaBat survey. Although we collected

one on the Site, it is unlikely that this

species regularly occurs there.

Reproduction

Parturition occurs in late May and

June (Lowery 1974, Schmidly 1991).

Litter size ranges from two to four

individuals and averages 3.4 per litter

(Jennings 1958). Mating probably

occurs in autumn and fertilization in

spring (Lowery 1974). The species’

reproductive habits in South Carolina

are not known.

Bats of South Carolina in Areas Neighboring the Savannah River Site
48 Bats of the Savannah River Site and Vicinity

Figure 42—Brazilian free-tailed bat.

Photo courtesy of John MacGregor

FAMILY MOLOSSIDAE

There are 16 genera and 86 species

in the family Molossidae (Nowak

1994). Members of this family are

found in southern Europe, Asia, Africa,

and from the northern half of South

America north into the United States.

Some are colonial and form the

largest aggregations of mammals

documented. Davis and others (1962)

estimated that summer molossid

populations in four caves (Bracken,

Goodrich, Rucker, and Frio) in Texas

exceeded 10 million individuals during

peak periods. Molossids are medium

to large bats. Because one of the most

defining characteristics of this family

is that the tail extends beyond the

posterior border of the uropatagium,

members of this family commonly are

called the free-tailed bats.

Two genera and six species of

molossids occur in the United States.

One species occurs in South Carolina.

Brazilian free-tailed bat

(Tadarida brasiliensis)

Morphology and Distribution

The Brazilian free-tailed bat is the

only species of free-tailed bat that

occurs in South Carolina (fig. 42).

As its name suggests, the tail of this

species extends beyond the posterior

edge of the uropatagium. It is the

smallest member of its genus in the

United States. Of the nine subspecies

now recognized, only T. brasiliensis

cynocephala occurs in the State

(Wilkins 1989). Its mass ranges from

8 to 14 g, and its average total length

is 91.9 mm (Whitaker and Hamilton

1998). See table 21 for average body

measurements of individuals collected

in the Southeast.

The Brazilian free-tailed bat has one

of the largest ranges of any mammal

in the Western Hemisphere (Wilkins

1989). From South Carolina, the

species’ range extends west into

southern Oregon and south through

Central America. It reaches the

southern limit of its range in the

Patagonian region of southern Chile

and Argentina (Wilkins 1989). A total

of 338 Brazilian free-tailed bats from

South Carolina are in museum

collections. Its distribution in the State

extends through the upper and lower

Coastal Plain (fig. 43). There have

been live captures in the Piedmont,

but these isolated records may not

reflect the species distribution in

South Carolina.

Free-tailed bats have been submitted

for rabies testing from Aiken and

Barnwell Counties, SC. Although none

was captured during the 1979 or 1996

to 2001 surveys, it was recorded

during a 1996 AnaBat study on

Savannah River Site (Menzel and

others 2002d). Additionally, its range

in the Southeast, the individuals

submitted for rabies testing from

Aiken and Barnwell Counties, and

recent reports that it is roosting in
49

Figure 43—Distribution of the Brazilian free-tailed bat in South Carolina (Menzel

and others 2003). Symbols denote counties for which records exist.

Table 21—Body measurements of the Brazilian free-tailed bat in the

Southeastern United States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -

Total 14 93.1 6.65 7.14 75.0 – 101.0

Tail 14 33.0 1.66 5.04 30.0 – 36.0

Foot 14 9.2 0.85 9.18 8.0 – 11.0

Ear 11 16.9 3.51 20.73 7.0 – 20.0

Tragus 7 1.7 0.64 37.11 1.0 – 3.0

Forearm 13 43.4 0.77 1.77 42.0 – 45.0

- - - - - - - - - - - - - - - - g - - - - - - - - - - - - - -

Mass 12 11.5 2.03 17.27 9.3 – 16.0

a Number of individuals measured.

structures near Clemson, SC, suggest

that it may be more common on the Site

than survey results indicate.6 Brazilian

free-tailed bats typically roost in humanmade structures. Their dependence on

such structures for roosting habitat may

limit their abundance on the Site.

Roosting

There is little difference between the

summer and winter roosting habits of

Brazilian free-tailed bats, although they

do vary among different regions of the

United States. Most roosts in the

Southwest are found in caves, although

the bat does use buildings sometimes

during migration. In the Southeast,

roosts typically are found in buildings

(Barbour and Davis 1969), but they

also have been found in hollow trees

and in the expansion joints of bridges

(Lowery 1974, Tuttle 1994). This

species is very gregarious and almost

always roosts in colonies (Barbour and

Davis 1969). Although the size of cave

colonies of the Southwest commonly

exceeds 1 million individuals, colonies

in the Southeast typically are

composed of < 50,000. Roosting habits

in South Carolina are not known.

Although nothing is known about the

species’ on-site roosting requirements,

there is no reason to suspect that

they are different from other areas

of the Southeast.

Foraging and Home Range

Although emergence time is

variable, Brazilian free-tailed bats

usually leave their roost shortly after

sunset (Bailey 1951). As with most bat

species inhabiting temperate regions,

the length of the foraging bout varies

with temperature. Bailey (1951) found

that on warm nights, individuals in a

colony of Brazilian free-tailed bats in

Louisiana did not return to the roost

until morning. Most studies of this

bat’s foraging habits have been

conducted in the Southwest. In the

Southeast, Brazilian free-tailed bats

feed on Hymenoptera, Coleoptera,

Diptera, and Lepidoptera (Sherman

6 Personal communication. 1999. Mary Bunch,

Wildlife Biologist, South Carolina Department of

Natural Resources, Wildlife Diversity Section,

P.O. Box 1806, Clemson, SC 29633.

Bats of South Carolina in Areas Neighboring the Savannah River Site
50 Bats of the Savannah River Site and Vicinity

Figure 44—Rocket-box bat house, as described by Dourson and MacGregor 1997.

Photo courtesy of John MacGregor

1939). Their diet in South Carolina

or on the Savannah River Site has not

been studied. There is no reason to

suspect that it would be any different

than elsewhere in the Southeast, but

because little is known about this

species in the region, and because

its diet may differ from the western

subspecies, its diet in the eastern

United States warrants future study.

Nothing is known about the size

of the Brazilian free-tailed bat’s home

range in the Southeast; further study

will be necessary to determine the

spatial extent of its foraging areas in

the region.

Effect of Habitat Type and

Stand Age on Flight Activity

We did not record any calls of

Brazilian free-tailed bats during the

AnaBat survey. However, the species

was detected during call surveys

conducted in canopy gaps in a

bottomland hardwood forest during

summer 1997 (Menzel and others

2002a). Brazilian free-tailed bats

have been collected from Aiken and

Barnwell Counties, SC, and from

areas south and north of the Site.

Reproduction

In Florida, spermatogenesis occurs

throughout late autumn and winter

(Sherman 1937). Unlike vespertilionid

bats in South Carolina, which mate in

autumn and ovulate in spring, Brazilian

free-tailed bats both copulate and

ovulate in spring (Cockrum 1955).

Breeding occurs in February and

March (Golley 1966). Parturition

occurs from late May to early August

(Barbour and Davis 1969, Sherman

1937), and the typical litter size is one

(Wilkins 1989). Maternity colonies

of this species have been located in

the State, and the species occasionally

may raise its young on the Savannah

River Site.

Conservation and

Research Needs

Although no federally listed bat

species occur on the Savannah

River Site, the State of South

Carolina lists the Rafinesque’s bigeared bat as endangered and the

southeastern myotis as threatened

(table 9). Two of the nine species

(22 percent) that occur on the Site

regularly are protected. In the

Southeast, 87 percent of all bat

species carry special conservation

designations somewhere within their

regional distribution (Laerm and

others 2000). Worldwide, a relatively

high proportion of bat species are

designated as threatened or

endangered. All 15 bat species

that occur in Great Britain are

now considered threatened or

endangered (Nagorsen and Brigham

1990, Stebbings 1988), and bat
51

populations seem to be declining

globally (Nowak 1994).

Potential issues of conservation

concern have been raised for the Site

and surrounding areas. More than 75

percent of the Site is included in the

timber management program and is

therefore subject to some timber

harvesting (Blake, in press). Although

there has been a recent surge of

information concerning the effects of

forest management practices on forest

bat communities, e.g., Barclay and

Brigham 1996, Krusic and others

1996, little is known about the effects

of most silvicultural practices in the

Southeast. Studies investigating the

effects that harvesting techniques,

e.g., clearcutting, group-selection cuts,

and various intermediate treatments,

e.g., thinnings, herbicide application,

and prescribed burning, may have on

bat communities will be necessary

to make more informed land

management decisions. As more

land area is devoted to short-rotation

forestry practices, roosting habitat

may become scarce for bat species

that depend on older trees for cavity or

bark roosts. Studies should investigate

the potential use of bat houses to

enhance roosting habitat where it

has been compromised (fig. 44).

In addition to timber management,

other large-scale agricultural practices

are common throughout much of

South Carolina. All nine species

of bats that occur on the Site are

insectivorous and therefore may be

affected by pesticides that are applied

to agricultural crops such as cotton,

soybeans, and corn. We know little

about how these pesticides will affect

bat populations over the long term.

The effects of pesticides on

cavernicolous species such as the

gray bat have been examined

(Clawson and Clark 1989), but the

effects of pesticides on tree-roosting

bats are not known. All bat species that

occur on the Savannah River Site roost

in trees during at least part of the year.

Another conservation concern in the

Coastal Plain of South Carolina focuses

on the use of bridges as roosts by bats.

In some areas of the United States,

such as Texas, bridges are important

roosting sites for many bat species

(Keeley 1997). The endangered

Rafinesque’s big-eared bat and the

threatened southeastern myotis roost

under bridges (Lance and others

2001). Although many bridges on the

Site may serve as roosts for these

species, no surveys have been

conducted to document the bats’

use of them.

In the 1990s, great strides were

made towards understanding the

roosting and foraging ecology of bats,

but much remains unknown. Few

studies have been conducted in the

Southeast, and results from studies

conducted in other regions of the

United States and Canada may not

be applicable in South Carolina or

on the Savannah River Site. To more

effectively manage the Site’s bat

communities we need more

information about their roosting

and foraging ecology.

Bats and Rabies

No work on bats is complete

without a discussion of rabies,

a zoonotic disease that

generates considerable concern

among public health officials and an

almost irrational fear among the

general public. Found worldwide

except in parts of Scandinavia, the

Australian continent, and on island

masses such as Hawaii, New Zealand,

Papua New Guinea, Japan, and Taiwan,

rabies is caused by exposure to and

infection by single-strand RNA viruses

in the genus Lyssavirus, family

Rhabdoviridae (Krebs and others

1995, Wunner and others 1988). The

British Isles had long been thought

to be rabies-free until the discovery

of a rabid Daubenton’s bat (Myotis

daubentonii) in 1996 (Whitby and

others 1996). Transmission to humans

or other mammals occurs primarily

by introduction of the virus into

wounds, cuts, scratches, and mucous

membranes from an infected animal,

usually by biting (Macinnes 1987).

Other routes of viral transmission,

such as inhalation of rabies-infected

aerosol, are known, but they are

extremely rare (Constantine 1962,

1967b; Winkler and others 1973), as is

transmission through rabies-infected

human tissue transplanted in

uninfected organ recipients (Centers

for Disease Control and Prevention

1980, 1981).

Rabies was first documented in bats

in the United States in 1953 (Krebs and

others 1994). Contrary to popular

belief, bats are not nonsymptomatic

carriers of rabies. However, the length

of latency prior to onset of symptoms

and eventual death in bats is highly

variable (Bell and others 1969,

Macinnes 1987, Moreno and Baer

1980). The rabies virus is reservoired

in many wild mammal hosts, including

bats (Davidson and Nettles 1997),

although transmission cycles usually

are separate between terrestrial

mammals and bats (Constantine 1967a,

1979a; Smith 1989). In Latin America,

vampire bats routinely spread rabies

to domestic livestock and humans

(Arellano-Sota 1988, Batista-da-Costa

and others 1993, Martinez-Burnes and

others 1997), and in North America,

cases of rabies in terrestrial wild

mammals and humans have resulted

from contact with several of the

insectivorous bat species commonly

found in South Carolina (Constantine

1979b). Previous research linked the

incidence of rabies in foxes to a

number of bat caves in the Karst

limestone regions of Tennessee

(Fredrickson and Thomas 1965),

but such relationships were poorly

substantiated (Fischman 1976).

Following the discovery of rabies virus

strains and species-specific variants,

most supposed links between bat

rabies and the maintenance of rabies

in terrestrial mammals have been

discounted (Krebs and others 1995,

Smith 1989, Smith and others 1986).

However, in areas where the incidence

of rabies in terrestrial mammals is low,

bats are responsible for small rabies

outbreaks or “clusters” (Daoust and

others 1996).

Among terrestrial mammals in South

Carolina and throughout the Southeast

and mid-Atlantic regions, raccoons

(Procyon lotor) are important

Conservation and Research Needs ° Bats and Rabies
52 Bats of the Savannah River Site and Vicinity

terrestrial reservoir species (Rosatte

and others 1997). Skunks (Memphitis

memphitis, Spilogale putorius) are

rabies vectors in the Midwest and

Mississippi Valley (Krebs and others

1994, 1995; Smith and others 1986),

and foxes (Urocyon cinereoargenteus,

Vulpes vulpes) are rabies reservoirs

in Canada, New England, and parts of

the Southwest (Davidson and Nettles

1997, Smith and others 1986). Coyotes

(Canis latrans) also appear to be

important vectors throughout much

of the Southwest (Clark and others

1994, Davidson and Nettles 1997).

In developing countries, human

cases of rabies are relatively common,

accounting for approximately 50,000

deaths annually (Fishbein and Bernard

1995). Most routes of human exposure

to rabies are from unvaccinated dogs

and cats, particularly in developing

countries, but wild animals, including

bats, are responsible for many human

cases (Krebs and others 1995).

Advances in public health—such as

animal control laws, mandatory

vaccination measures for dogs in many

jurisdictions, and aggressive and

prompt medical attention following

animal bites—have served to limit

human exposure to and infection with

rabies in North America (Rosatte and

others 1997, Tierkel 1975). Today,

human exposure to rabies in the

United States is almost wholly limited

to contact from wild animals, of which

exposure to infected bats accounts for

the majority of cases documented

since the 1980s (Childs and others

1994, Hoff and others 1993, Krebs and

others 1995). Nonetheless, the overall

incidence of rabies contracted from

bats in North America is low (Krebs

and others 1995, Rotz and others 1998,

Tuttle and Kern 1981). Efforts to

develop effective oral rabies vaccines

for wild, terrestrial mammals—and to

distribute such vaccines by widespread

baiting—are underway in the United

States and Canada (Davidson and

Nettles 1997), but such preventive

measures for bats have not been

contemplated.

In humans, rabies is almost always

fatal. Following exposure, the

incubation period may range from

< 10 days to > 6 years (extreme

latency). Commonly, incubation time

from exposure to the first symptoms is

30 to 90 days. The rabies virus infects

nervous tissue, eventually spreading to

the spinal cord and brain, producing

encephalitic lesions. It moves then

throughout the central nervous system

and to peripheral sites such as the

salivary glands, where it is shed in

saliva (thus facilitating transfer to

another animal). Initial symptoms in

humans can include pain and burning

near the site of the wound where

exposure occurred, headache, fever,

flu-like malaise, and mental changes

such as anxiety and apprehension. As

the disease progresses, mental

deterioration occurs in the form of

disorientation and hallucinations.

Aggressive behavior and hydrophobia,

or “fear of water,” does occur in human

cases, although in less than one-half of

documented rabies cases. Physically,

difficulty in swallowing, muscle

spasms, and paralysis occur as the

disease advances; death usually results

from respiratory failure. From the

onset of initial symptoms, death may

occur within 1 to a few weeks

(Fishbein 1991, Jackson and others

2003, Krebs and others 1995).

In wild mammals, rabies takes two

forms—a furious or aggressive form

and a withdrawn or dumb form.

Skunks, foxes, dogs, and raccoons

often lose all fear of humans and other

animals when exhibiting the furious

form of the disease, and they may

exhibit extremely aggressive

behaviors. Activity patterns may

change. For example, nocturnal

animals may become active during

the day (Davidson and Nettles 1997,

Macinnes 1987). Rabid bats may be

irritable and bite conspecifics, but

furious behaviors, such as are seen in

wild carnivores, are rare (Macinnes

1987, Sulkin and Allen 1974, Tuttle and

Kern 1981). The extent to which bat-tobat rabies transmission occurs through

contact from grooming and nursing or

from antagonistic interspecific

encounters is unknown (Krebs and

others 1995, Macinnes 1987). Actual

unprovoked attacks by rabid,

insectivorous bats on humans or

other animals are extremely rare

(Constantine 1979a, 1979b; Tuttle and

Kern 1981). Animals infected with

rabies from bats often exhibit a

creeping paralysis (Baer 1975). In the

dumb form of rabies, animals remain

calm, yet are apparently disoriented

and confused (Davidson and Nettles

1997, Macinnes 1987). In most wild

mammals, symptoms similar to those

affecting humans occur as the disease

progresses, advancing to paralysis and

eventual death.

Following a suspected or confirmed

exposure to rabies—via the use of a

fluorescent antibody test on brain

tissue of the suspect animal (Kissling

1975, Wachendorfer and others

1985)—development of rabies can be

prevented by prompt postexposure

prophylaxis of human antirabies

immunoglobulin injections. Such

treatment is followed by a series of

human diploid cell injections, rabiesadsorbed intramuscular vaccinations,

or purified chick embryo cell culture

vaccinations (Blythe and others 1998).

For veterinarians, animal control

officers, wildlife biologists, wildlife

rehabilitators, and zoologists, who

come into contact with wild and feral

mammals on a regular basis,

preexposure prophylaxis is

recommended (Krebs and others

1995). Similarly, anyone handling bats

or coming in close contact with bats

should be preimmunized.

For many years, rabies was thought

to be a single viral type. With the

advent of monoclonal antibody surveys

and advances in genetics research

through polymerase chain reaction

(PCR) techniques, epidemiologists

have documented a number of

antigenically and genetically distinct

variants or strains of rabies virus

(Koprowski and others 1985; Smith

and others 1984, 1986). Strains unique

to several species of bats throughout

North America have been documented

(Smith 1988), and within a species

there may be more than one strain or

variant across wide geographic regions

or within a localized area (Smith and

others 1984). Thus, rabies strains
53

found in bats are distinct from those

in terrestrial mammals. As a result,

public health officials and scientists

can differentiate and document routes

of human exposure by animal type

and species. For example, of the 14

confirmed cases of human rabies

in the United States contracted

domestically from 1980 to 1994, 11

were linked to bats. Of those, eight

were attributed to viral strains

associated with silver-haired bats

(Krebs and others 1995), although that

species was not always the confirmed

vector (Morimoto and others 1996),

nor was evidence of a bat bite always

present. Common carriers of the silverhaired bat rabies viral strain appear to

be big brown bats, little brown bats,

and eastern pipistrelles (Messenger

and others 1997).

Rabies has been documented in

most bat species that occur in South

Carolina, including hoary bat, red bat,

yellow bat, Seminole bat, big brown

bat, little brown bat, small-footed bat,

southeastern bat, gray bat, Townsend’s

big-eared bat, eastern pipistrelle,

evening bat, silver-haired bat, and

Brazilian free-tailed bat. Rabies has not

been documented in Rafinesque’s bigeared bat (Constantine 1979b). The

common species most likely to occur

in and near human dwellings and

activity centers, such as the big brown

bat, probably pose the greatest risk for

transmitting rabies to humans (Childs

and others 1994). Eastern red bats,

perhaps the most widespread and

numerous species in South Carolina,

accounted for a large percentage of

rabid bats collected in Illinois (Burnett

1989), while a high percentage of

rabies-positive bats in Florida have

been yellow bats (Bigler and others

1975). Nationwide, the Centers for

Disease Control and Prevention

statistics indicate that approximately

10 percent of all annually reported and

confirmed rabid animals are bats

(Krebs and others 1995). For example,

in 1996, there were 7,124 documented

cases of nonhuman rabies in the

United States, of which 741 were bats

(Krebs and others 1997). Still, the

overall human health risks posed by

rabid bats are extremely small in South

Carolina. Simple preventive measures

such as avoiding handling any bat and

avoiding entry into caves, attics, and

abandoned buildings that harbor

roosting bats—or preventing roosting

bats from using human-occupied

dwellings—would eliminate most

routes of contact and potential rabies

transmission. Immediate postexposure

rabies prophylaxis should proceed if

one is bitten by or comes into close

contact with a bat, such as waking

to find a bat present in occupied

sleeping quarters.

Acknowledgments

We thank the U.S. Department

of Energy; the USDA Forest

Service-Savannah River,

Southern Research Station, and

Northeastern Research Station; the

National Council of the Paper Industry

for Air and Stream Improvement; West

Virginia University Division of Forestry;

and West Virginia University Arlen G.

and Louise Stone Swiger Fellowship for

supporting this project. Funding from

the U.S. Department of Energy was

provided through the USDA Forest

Service-Savannah River under

Interagency Agreement DE-IA09-

76SR00056. This project would not

have been possible without the

professional support of Drs. B.R.

Chapman and J. Laerm. Drs. B.R.

Chapman and S.B. Castleberry provided

helpful comments on

the manuscript. This work was

conceived and supported throughout

by Dr. J.I. Blake.

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61

Table 22—Dental formulas for the bats of the Savannah River Site

Upper teetha Lower teetha Total

Species I C Pm M I C Pm M (x2)

Eastern pipistrelle 2 1 2 3 3 1 2 3 34

Southeastern myotis 2 1 3 3 3 1 3 3 38

Little brown bat 2 1 3 3 3 1 3 3 38

Evening bat 1 1 1 3 3 1 2 3 30

Rafinesque’s big-eared bat 2 1 2 3 3 1 3 3 36

Silver-haired bat 2 1 2 3 3 1 3 3 36

Eastern red bat 1 1 2 3 3 1 2 3 32

Seminole bat 1 1 2 3 3 1 2 3 32

Northern yellow bat 1 1 1 3 3 1 2 3 30

Hoary bat 1 1 2 3 3 1 2 3 32

Big brown bat 2 1 1 3 3 1 2 3 32

Brazilian free-tailed batb 1 1 2 3 2/3 1 2 3 30/32

I = incisors; C = canines; Pm = premolars; M = molars.

a Number of teeth in each side of jaw.

b Although the Brazilian free-tailed bat has not been captured on the Savannah River Site, it is likely this species occasionally

occurs there.

Appendix

Appendix

KEYS TO THE SKINS

AND SKULLS OF SOUTH

CAROLINA BATS

Bat research as well as bat

monitoring in the Southeast

have been impeded by

difficulty in training field personnel

to reliably identify captured bats.

Although many published keys contain

information about particular species in

the region (Barbour and Davis 1969,

Golley 1962, Hoffmeister 1989, Jenkins

1949, Schmidly 1991, Whitaker and

Hamilton 1998), many of them have

been unreliable tools for teaching

students and field personnel to identify

bats by their skins and skulls. Most

keys require the lower jaw, are poorly

or inadequately illustrated, or contain

unreliable univariate discriminatory

measurements. To improve field

identification capabilities we devised a

tool combining traits that have been

useful in the field and the lab with

information from other published

keys. Using the combined keys has

facilitated relatively easy and accurate

identification of the 14 bat species

that occur in South Carolina. The

keys are reprinted from Menzel and

others (2002a).

We used measurements from

museum specimens, information

contained in recent publications about

bat morphological characteristics,

and information from published

dichotomous keys (Barbour and Davis

1969, Golley 1962, Hoffmeister 1989,

Jenkins 1949, Schmidly 1991). Because

the bat’s lower jaw often is damaged

or missing from museum skeletal

specimens or skulls found in the field,

we devised a key that does not require

the lower jaw for identification.

The eastern red (Lasiurus borealis)

and Seminole (L. seminolus) bats are

two of the most common species on

the Site, but many keys to bats in the

Southeast do not differentiate between

their skulls. Because these two species

are relatively abundant on the Site, we

were able to incorporate information

about the size of the protuberance of

the lacrimal ridge, i.e., the lacrimal

shelf, and accurately differentiate

among about 75 percent of eastern

red and Seminole bats skulls found

there (Laerm and others 1999,

Lowery 1974).

The key to the skulls allows a user

to classify a specimen as belonging to

the genus Myotis. Because simple

univariate measures cannot be used

to differentiate among southeastern

myotid skulls, identification to species

cannot be accomplished with this key.

However, many nonmyotid bat skulls

can be identified to species by

counting the number of teeth in one

upper quadrant (one-half of the upper

jaw), measuring the skull’s greatest

length (from the posteriormost margin

to the anteriormost portion, not

including the incisors), and comparing

these lengths using the dental formula

for each species (table 22) and the

skull key.
62 Bats of the Savannah River Site and Vicinity

Figure 45—Location of the standard body measurements on a bat, including total length (TL), tail length (TV), forearm length

(FA), and foot length (HF). Inset illustrates the extent of fusion in the epiphyseal gap of the finger joints of adults and juveniles.

Adult joints (AD) appear fused and consist of a single protuberance; juvenile joints (JV) are not fused (cartilaginous plates

remain in the joints) and consist of two protuberances or a single protuberance that is larger and more tapered than in adults. Illustration courtesy of B. Coslick
63

Figure 46—Location of ear (E) and tragus length

(TR) measurements.

Illustration courtesy of B. Coslick

Appendix

Locations of the six standard

body measurements—total length

(TL), tail length (TV), foot length

(HF), ear length (E), forearm length

(FA), and tragus length (TR)—are

illustrated in figures 45 and 46. These

measurements can be taken from dead

specimens prior to preparation or from

live specimens prior to release.

Because we designed the keys using

characteristics and measurements

taken from adult individuals, they

may not be reliable when identifying

juveniles. Juvenile and adult age

classes can be determined by

examining the extent of epiphysealdiaphyseal fusion in the finger joints

(Anthony 1988). Cartilaginous plates

may not be apparent in the finger joints

of adults; the joints may consist of a

single, knobby protuberance (fig. 45).

Cartilaginous plates are apparent in

the finger joints of juveniles; the joints

consist of two protuberances with a

slight taper between or a single

protuberance that is much longer

and more tapered at both ends than

the protuberance in adult joints (fig.

45). The most effective way to observe

the cartilaginous plates is by backlighting the wing and looking for

semitransparent sections in joints

of the phalanges. We also have

included a summary table listing

the characteristics of each species

(table 23).
64 Bats of the Savannah River Site and Vicinity Table 23—A comparison of characteristics of bats of the Savannah River Site Species Pelage Forearm Weight Foot Ear Tragus shapea Calcara Other identifying traits mm g mm Brazilian free-tailed bat (Tadarida brasiliensis) Short, brown to dark gray 43 11–14 8–11 19–20 Tail free from membrane Rafinesque’s big-eared bat (Corynorhinus rafinesquii) Gray or brown above and whitish below 41 7–10 8–13 27–37 Broad and long Large ears Eastern red bat Bright red or rust color, females with frosting, Short, blunt, Membrane between rear legs is (Lasiurus borealis) males without 40 8–16 9–10 17–18 and curved completely furred Hoary bat Yellowish brown to dark brown with very heavy Short, blunt, Very large; membrane between (Lasiurus cinereus) frosted tips 53 20–35 6–13 17 and curved rear legs is completely furred

Eastern pipistrelle Blunt and straight Not(Pipistrellus subflavus) Sandy brown to orange on back—tricolored 33 4–7 8–10 13–15 keeled Pink forearms, black wings

Silver-haired bat Blunt and Membrane between rear legs is(Lasionycteris noctivagans) Dark blackish brown, with silver frosted tips 41 7–16 7–11 14 rounded furred on anterior halfBig brown bat Broad and Large; prominent glands(Eptesicus fuscus) Long brown, with two-toned black base 47 13–25 10–12 17–18 rounded Keeled on mussel

Evening bat Not Strong odor, prominent glands(Nycteceius humeralis) Dull brown with paler belly, young much darker 36 5–14 7–8 14–15 Short and rounded keeled on mussel, single upper incisorEastern small-footed myotis Sharply Black mask, relatively long tail(Myotis leibii) Blackish brown above, paler below 32 4–6 6–8 13–15 Sharp keeled (33 mm)

Southeastern myotis Wooly, dull, little contrast between base and tips, Slender and Not Gray-black wings, long(Myotis austroriparius) yellowish to gray above, white below 40 5–12 10–13 13–16 pointed keeled toe hairsNorthern long-eared myotis Very long and Large propatagium, ears(Myotis septentrionalis) Not glossy, brown above, grayish below 35 5–10 7–10 16–19 slender Keeled extend beyond nose (3 mm)

Little brown bat Long glossy tan to dark brown above, gray to Slender and Not Long toe hairs (extending beyond(Myotis lucifugus) buff below 38 7–8 8–11 13–16 pointed keeled claws), wings contrasting with bodya Blank cells indicate characteristics not important for species identification.
65

Figure 47—Contrast between the tail and uropatagium of the Brazilian free-tailed

bat and the other 15 species of bats that occur in Georgia. The (A) tail of the

Brazilian free-tailed bat extends beyond the posterior margin of the uropatagium;

(B) tails of the other 15 bat species are completely enclosed in the uropatagium.

Illustration courtesy of B. Coslick

Appendix

Key to the Skins

1. a. One-third or more of the tail

extending beyond uropatagium

(fig. 47a)—Brazilian free-tailed

bat (Tadarida brasiliensis)

b. Tail not extending beyond

uropatagium, or only slightly

(fig. 47b)—2

2. a. Dorsal surface of uropatagium

at least partially furred—3

b. Dorsal surface of uropatagium

not furred or slightly furred at

the junction with the body—7

3. a. Pelage black, tips of hairs

frosted with white—4

b. Pelage red, mahogany, or

yellow—5

4. a. Total length > 120 mm,

uropatagium heavily furred

throughout, ear white or yellow

with black rim—hoary bat

(Lasiurus cinereus)

b. Total length > 115 mm, posterior

one-third of uropatagium bare, ear

solid black—silver-haired bat

(Lasionycteris noctivagans)

5. a. White shoulder patch absent,

yellow coloration, frosting

absent—northern yellow bat

(Lasiurus intermedius)

b. White shoulder patch present,

red or mahogany coloration,

frosting usually present (except

male Lasiurus borealis)—6

6. a. Pelage bright-brick red, tips of

hair frosted white (except males),

face light red/yellow—eastern red

bat (Lasiurus borealis)

b. Pelage dark mahogany, tips

of hair frosted white, face

mahogany/red—Seminole bat

(Lasiurus seminolus)
66 Bats of the Savannah River Site and Vicinity

Figure 48—Large ears and location of the pararhinal glands on Rafinesque’s big-eared bat.

Figure 49—Comparison of (A) the short, blunt tragus characteristic of species such as the eastern

pipistrelle and evening bat; and (B) the long, pointed, sharp tragus characteristic of Myotis.

Illustration courtesy of B. Coslick

Illustration courtesy of B. Coslick
67

Figure 50—Comparison of a (A) keeled calcar characteristic of smallfooted myotis and (B) an unkeeled calcar characteristic of little brown

and southeastern myotis.

Illustration courtesy of B. Coslick

Appendix

7. a. Ears > 25 mm long, distinctive

pararhinal glands (large bumps,

fig. 48) on either side of nose—

Rafinesque’s big-eared bat

(Corynorhinus rafinesquii)

b. Ears < 25 mm long, pararhinal

glands not distinct—8

8. a. Total length > 100 mm,

forearm > 40 mm—big brown

bat (Eptesicus fuscus)

b. Total length < 100 mm,

forearm < 40 mm—9

9. a. Tragus (projection within

the ear) short, blunt, and curved

(fig. 49a)—10

b. Tragus long, pointed at tip,

and straight (fig. 49b)—11

10. a. Dorsal fur tricolored when

parted; coloration black at base,

yellowish brown in the middle and

dark brown at tips; forearm pink

and < 32 mm—eastern pipistrelle

(Pipistrellus subflavus)

b. Dorsal fur dark brown, forearm

dark and >32 mm long—evening

bat (Nycticeius humeralis)

11. a. Ear > 16 mm long, extends >2

mm beyond the tip of nose when

laid forward—northern long-eared

myotis (Myotis septentrionalis)

b. Ear does not extend beyond the

tip of nose when laid forward—12

12. a. Calcar keeled (figs. 50a and

51a)—eastern small-footed myotis

(Myotis leibii)

b. Calcar not keeled (figs. 50b

and 51b)—13

13. a. Tips of hairs reddish; hair long

and glossy—little brown bat

(Myotis lucifugus)

b. Tips of hairs not reddish; hair

short and wooly—southeastern

myotis (Myotis austroriparius)
68 Bats of the Savannah River Site and Vicinity

Figure 51—Photograph comparing (A) the keeled calcar characteristic of small-footed myotis to

(B) an unkeeled calcar characteristic of little brown and southeastern myotis.

A

B
69

Figure 52—Comparison of (A) the

upper incisors of Brazilian free-tailed

bats, and (B) upper incisors of the

other southeastern bats. Upper

incisors of the Brazilian free-tailed bat

converge at the tips; upper incisors of

other southeastern bats do not.

Figure 53—Comparison of the

lacrimal ridge protuberance of (A)

eastern red and (B) Seminole bats.

Illustrations depict a dorsal surface

view of the upper left skull quadrant.

Note that the two triangular

structures at the top are the upper

incisor and the upper canine. The

hooked structure at the lower left

section is the anterior-most section

of the zygomatic arch. The lacrimal

ridge protuberance (lacrimal shelf)

typically is larger in (A) eastern red

bats than in (B) Seminole bats.

Illustration courtesy of B. Coslick

Illustration courtesy of G. Weis-Gresham

Appendix

Key to Skulls

1. a. Upper incisors obviously

converge at tips, i.e., much closer

together at tips than at base (fig.

52a—Brazilian free-tailed bat

(Tadarida brasiliensis)

b. Upper incisors wider at tip than

base, equidistant at tip and base, or

tips slightly converging (fig. 52b);

premaxillary gap square—2

2. a. Nine teeth present in upper

quadrant—Myotis spp.

b. Fewer than nine teeth in upper

quadrant—3

3. a. Eight teeth in upper quadrant—4

b. Fewer than eight teeth in upper

quadrant—6

4. a. Upper incisor bifid (twocusped)—Rafinesque’s big-eared

bat (Corynorhinus rafinesquii)

b. Upper incisor unicuspid—5

5. a. Greatest length of skull

> 13.5 mm; rostrum flat

with two concavities on dorsal

surface—silver-haired bat

(Lasionycteris noctivagans)

b. Greatest length of skull

< 13.5 mm, rostrum sloped

with no concavities on dorsal

surface—eastern pipistrelle

(Pipistrellus subflavus)

6. a. Seven teeth in upper quadrant—7

b. Six teeth in upper quadrant—10

7. a. Two upper incisors (one large,

one minute)—big brown bat

(Eptesicus fuscus)

b. One upper incisor—8

8. a. Greatest skull length >15.5

mm—hoary bat (Lasiurus cinereus)

b. Greatest skull length <15.5

mm—9

9. a. Protuberance of the lacrimal

ridge (shelf) well developed

(fig. 53a)—eastern red bat

(Lasiurus borealis)

b. Protuberance of the lacrimal

ridge poorly developed or

absent (fig. 53b)—Seminole bat

(Lasiurus seminolus)

10. a. Sagittal crest well developed,

greatest length of skull

> 16 mm—northern yellow bat

(Lasiurus intermedius)

b. Sagittal crest absent or poorly

developed, greatest length of

skull < 16 mm—evening bat

(Nycticeius humeralis)
70 Bats of the Savannah River Site and Vicinity
Menzel, Michael A.; Menzel, Jennifer M.; Kilgo, John C. [and others]. 2003. Bats

of the Savannah River Site and vicinity. Gen. Tech. Rep. SRS-68. Asheville, NC: U.S.

Department of Agriculture, Forest Service, Southern Research Station. 69 p.

The U.S. Department of Energy’s Savannah River Site supports a diverse bat community.

Nine species occur there regularly, including the eastern pipistrelle (Pipistrellus

subflavus), southeastern myotis (Myotis austroriparius), evening bat (Nycticeius

humeralis), Rafinesque’s big-eared bat (Corynorhinus rafinesquii), silver-haired bat

(Lasionycteris noctivagans), eastern red bat (Lasiurus borealis), Seminole bat (L.

seminolus), hoary bat (L. cinereus), and big brown bat (Eptesicus fuscus). There are

extralimital capture records for two additional species: little brown bat (M. lucifigus) and

northern yellow bat (Lasiurus intermedius). Acoustical sampling has documented the

presence of Brazilian free-tailed bats (Tadarida brasiliensis), but none has been captured.

Among those species common to the Site, the southeastern myotis and Rafinesque’s bigeared bat are listed in South Carolina as threatened and endangered, respectively. The

presence of those two species, and a growing concern for the conservation of forest-dwelling

bats, led to extensive and focused research on the Savannah River Site between 1996 and

2002. Summarizing this and other bat research, we provide species accounts that discuss

morphology and distribution, roosting and foraging behaviors, home range characteristics,

habitat relations, and reproductive biology. We also present information on conservation

needs and rabies issues; and, finally, identification keys that may be useful wherever the bat

species we describe are found.

Keywords: Bats, foraging, habitat use, rabies, roosting, Savannah River Site.
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resources for sustained yields of wood, water, forage, wildlife, and

recreation. Through forestry research, cooperation with the States

and private forest owners, and management of the National Forests

and National Grasslands, it strives—as directed by Congress—

to provide increasingly greater service to a growing Nation.

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call 202-720-5964 (voice and TDD). USDA is an equal

opportunity employer.

A publication of the Savannah River Site National

Environmental Research Park Program. Publication

number: SRO-NERP-26