united states agriculture bats of the savannah river site ... · savannah river site and vicinity...

United StatesDepartment ofAgriculture

Forest Service

SouthernResearch Station

General TechnicalReport SRS-68

Bats of theSavannah RiverSite and VicinityMichael 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

1Graduate Research Assistant, Division of Forestry, Wildlife andFisheries, West Virginia University, Morgantown, WV 26506;2Research Wildlife Biologist, Northeastern Research Station,USDA Forest Service, Parsons, WV 26287; 3Research Wildlife

Biologist, Southern Research Station, USDA Forest Service, NewEllenton, SC 29809; 4Graduate Research Assistant, Department ofZoology, Southern Illinois University, Carbondale, IL 62901; and5Assistant 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 JohnMacGregor); Rafinesque’s big-eared bat (photo by John

MacGregor); eastern red bat (photo by John MacGregor); andeastern red bat (photo by Julie Roberge).

September 2003

Southern Research StationP.O. Box 2680

Asheville, NC 28802

i

Bats of theSavannah RiverSite and VicinityMichael A. Menzel, Jennifer M. Menzel, John C. Kilgo,W. Mark Ford, Timothy C. Carter, and John W. Edwards

AbstractThe U.S. Department of Energy’s Savannah River Site supports a diversebat community. Nine species occur there regularly, including the easternpipistrelle (Pipistrellus subflavus), southeastern myotis (Myotis austroriparius),evening bat (Nycticeius humeralis), Rafinesque’s big-eared bat (Corynorhinusrafinesquii), silver-haired bat (Lasionycteris noctivagans), eastern red bat(Lasiurus borealis), Seminole bat (L. seminolus), hoary bat (L. cinereus), andbig brown bat (Eptesicus fuscus). There are extralimital capture records fortwo additional species: little brown bat (M. lucifigus) and northern yellow bat(Lasiurus intermedius). Acoustical sampling has documented the presence ofBrazilian free-tailed bats (Tadarida brasiliensis), but none has been captured.Among those species common to the Site, the southeastern myotis andRafinesque’s big-eared bat are listed in South Carolina as threatened andendangered, respectively. The presence of those two species, and a growingconcern for the conservation of forest-dwelling bats, led to extensive andfocused research on the Savannah River Site between 1996 and 2002.Summarizing this and other bat research, we provide species accounts thatdiscuss morphology and distribution, roosting and foraging behaviors, homerange characteristics, habitat relations, and reproductive biology. We alsopresent information on conservation needs and rabies issues; and, finally,identification keys that may be useful wherever the bat species we describeare found.

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

A publication of the Savannah River Site National Environmental ResearchPark Program. Publication number: SRO-NERP-26

iii

Table of Contents

1 Introduction2 Major Vegetation Types of

the Savannah River Site6 The Sitewide Bat Survey

13 Accounts of Species14 Bats of the Savannah River Site14 FAMILY VESPERTILIONIDAE

14 Eastern pipistrelle (Pipistrellus subflavus)Morphology and Distribution .............................................................. 14Roosting ................................................................................................. 14Foraging and Home Range .................................................................. 16Effect of Habitat Type and Stand Age on Flight Activity .................. 16Reproduction ......................................................................................... 16

18 Southeastern myotis (Myotis austroriparius)Morphology and Distribution .............................................................. 18Roosting ................................................................................................. 19Foraging and Home Range .................................................................. 19Effect of Habitat Type and Stand Age on Flight Activity .................. 19Reproduction ......................................................................................... 21

21 Evening bat (Nyticeius humeralis)Morphology and Distribution .............................................................. 21Roosting ................................................................................................. 23Foraging and Home Range .................................................................. 23Effect of Habitat Type and Stand Age on Flight Activity .................. 23Reproduction ......................................................................................... 23

25 Rafinesque’s big-eared bat (Corynorhinus rafinesquii)Morphology and Distribution .............................................................. 25Roosting ................................................................................................. 26Foraging and Home Range .................................................................. 26Effect of Habitat Type and Stand Age on Flight Activity .................. 26Reproduction ......................................................................................... 26

27 Silver-haired bat (Lasionycteris noctivagans)Morphology and Distribution .............................................................. 27Roosting ................................................................................................. 27Foraging and Home Range .................................................................. 27Effect of Habitat Type and Stand Age on Flight Activity .................. 28Reproduction ......................................................................................... 28

iv

Contents, cont.

29 Eastern red bat (Lasiurus borealis)Morphology and Distribution .............................................................. 29Roosting ................................................................................................. 29Foraging and Home Range .................................................................. 30Effect of Habitat Type and Stand Age on Flight Activity .................. 30Reproduction ......................................................................................... 32

32 Seminole bat (Lasiurus seminolus)Morphology and Distribution .............................................................. 32Roosting ................................................................................................. 33Foraging and Home Range .................................................................. 34Effect of Habitat Type and Stand Age on Flight Activity .................. 34Reproduction ......................................................................................... 34

34 Hoary bat (Lasiurus cinereus)Morphology and Distribution .............................................................. 34Roosting ................................................................................................. 35Foraging and Home Range .................................................................. 35Effect of Habitat Type and Stand Age on Flight Activity .................. 37Reproduction ......................................................................................... 37

37 Big brown bat (Eptesicus fuscus)Morphology and Distribution .............................................................. 37Roosting ................................................................................................. 39Foraging and Home Range .................................................................. 39Effect of Habitat Type and Stand Age on Flight Activity .................. 39Reproduction ......................................................................................... 39

41 Bats of South Carolina in AreasNeighboring the Savannah River Site

41 FAMILY VESPERTILIONIDAE

41 Little brown bat (Myotis lucifugus)Morphology and Distribution .............................................................. 41Roosting ................................................................................................. 42Foraging and Home Range .................................................................. 42Effect of Habitat Type and Stand Age on Flight Activity .................. 42Reproduction ......................................................................................... 42

43 Eastern small-footed myotis (Myotis leibii)Morphology and Distribution .............................................................. 43Roosting ................................................................................................. 44Foraging and Home Range .................................................................. 44Effect of Habitat Type and Stand Age on Flight Activity .................. 44Reproduction ......................................................................................... 44

v

Contents, cont.

44 Northern long-eared myotis (Myotis septentrionalis)Morphology and Distribution .............................................................. 44Roosting ................................................................................................. 45Foraging and Home Range .................................................................. 45Effect of Stand Age and Habitat Type on Flight Activity .................. 45Reproduction ......................................................................................... 45

46 Northern yellow bat (Lasiurus intermedius)Morphology and Distribution .............................................................. 46Roosting ................................................................................................. 46Foraging and Home Range .................................................................. 46Effect of Habitat Type and Stand Age on Flight Activity .................. 47Reproduction ......................................................................................... 47

48 FAMILY MOLOSSIDAE

48 Brazilian free-tailed bat (Tadarida brasiliensis)Morphology and Distribution .............................................................. 48Roosting ................................................................................................. 49Foraging and Home Range .................................................................. 49Effect of Habitat Type and Stand Age on Flight Activity .................. 50Reproduction ......................................................................................... 50

50 Conservation and Research Needs51 Bats and Rabies53 Acknowledgments53 Literature Cited61 Appendix61 KEYS TO THE SKINS AND SKULLS

OF SOUTH CAROLINA BATS

Key to the Skins .................................................................................... 65Key to Skulls .......................................................................................... 69

1

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

Wing Wing-aspect Wing loadingSpecies Mass length ratio indexa indexb

g mm

Eastern pipistrelle 5.4 97.7 2.27 1.28Southeastern myotis 5.7 115.0 2.30 0.99Little brown bat 6.5 107.7 2.23 1.26Evening bat 8.0 114.0 2.43 1.49Rafinesque’s big-eared bat 8.6 127.0 2.27 1.21Silver-haired bat 9.3 128.3 2.57 1.45Eastern red bat 10.0 138.0 2.64 1.39Seminole bat 10.1 138.3 2.66 1.59Northern yellow bat 20.0 158.3 2.58 2.06Hoary bat 21.2 183.0 2.94 1.88Big brown bat 15.3 138.3 2.47 1.97Brazilian 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 speciesoccasionally occurs onsite.

Table 1—Comparison of average mass, total length, and forearm length ofbats of the Savannah River Site

Species Mass Total length Forearm

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

Eastern pipistrelle 5.4 fg 83.6 h 34.3 gSoutheastern myotis 5.7 efg 86.6 gh 37.1 fgLittle brown bat 6.5 defg 87.4 gh 37.0 fgEvening bat 8.0 cdefg 86.8 gh 36.2 fgRafinesque’s big-eared bat 8.6 cdef 96.1 ef 43.4 cdSilver-haired bat 9.3 cde 99.5 def 41.6 deEastern red bat 10.0 cd 101.8 de 39.5 efSeminole bat 10.1 c 103.9 d 41.5 deNorthern yellow bat 20.0 a 127.6 b 51.7 bHoary bat 21.2 a 132.5 a 55.2 aBig brown bat 15.3 b 111.6 c 45.9 cBrazilian 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 thisspecies occasionally occurs onsite.

Introduction

T he Savannah River Sitesupports a diverse batcommunity. Nine species occur

there regularly. Another three mayuse the Site occasionally. These three,the northern yellow bat (Lasiurusintermedius Allen), little brown bat(Myotis lucifugus LeConte), andBrazilian free-tailed bat (Tadaridabrasiliensis I. Geof. St.-Hilaire), havebeen documented on the Site butprobably occur there only occasionally.

Relative to other mammalian orders,the size of these 12 species varies little.Total body length ranges from 84 mmto 132 mm, and wing length rangesfrom 98 mm to 183 mm (see tables 1and 2 and tables accompanying eachspecies account). In terms of mass,the largest bat known to occur on theSite, the hoary bat (Lasiurus cinereusBeauvois), is only four times largerthan the smallest bat found there,the eastern pipistrelle (Pipistrellussubflavus F. Cuvier) (table 1). Bycomparison, the largest rodent there,the beaver (Castor canadensis Kuhl),is 200 times larger than the smallestrodent, the eastern harvest mouse(Reithrodontomys humulis Audubonand Bachman), and the largestcarnivore, the black bear (Ursusamericanus Palla), is more than900 times larger than the smallestcarnivore, the long-tailed weasel(Mustela frenata Lichtenstein)(Cothran and others 1991).

Distributions of most temperatezone bats are strongly influencedby the availability of roost structures(Findley 1993, Humphrey 1975).The roosting habits of bats in SouthCarolina are diverse (table 3), resultingin variations in the diversity of batcommunities across the State. Theleast diverse bat community occurs inthe Piedmont (6 species), whereas themost diverse occur in the Blue Ridgeand lower Coastal Plain (12 species)(Menzel and others 2003). Nine of thefourteen species that occur in SouthCarolina (Menzel and others 2003)typically roost in caves or mines during

Bats of the Savannah River Site and Vicinity2

part of the year (table 3). Threespecies roost in mines or caves, butonly during the winter (table 3).In addition to suitable foliage andcavity roost sites, the Blue Ridge alsocontains mines and crevices in verticalrock faces, resulting in the greatestdiversity of roost structures in theState. Thus diversity in the batcommunity on the Savannah RiverSite may be somewhat limited simplyby its physiographic position.

In the last 10 years many studieshave focused on the distributionand natural history of bats in theSoutheastern United States. Overhalf the total published literatureabout the natural history of bats inSouth Carolina and Georgia waspublished in that period, and researchhas proceeded as fast in otherSoutheastern States. Although manyaspects of their behavior and naturalhistory in the Southeast remain poorlyunderstood, much is known about

bat species’ roosting requirements,foraging behavior, diet, home range,and reproduction. Making landmanagement decisions without regardfor the effects they may have on batcommunities no longer can be justifiedby claims that too little is known abouthabitat requirements of this diversemammalian order.

Many recent studies of bat habitatrequirements were conducted on theSite. Similar studies elsewhere inthe Southeast also provided usefulinformation about the natural historyof bats known to occur there. Thisreport summarizes information aboutthe distribution, roosting ecology,foraging behavior, diet, home range,and reproduction of bats that occuron and near the Site. Informationcontained herein should aid landmanagers in assessing how theirmanagement decisions may affectbat species.

Major VegetationTypes of the

Savannah River Site

T he Savannah River Site isa former nuclear weaponsmaterial production facility

operated by the U.S. Departmentof Energy (DOE). It borders theSavannah River in west-central SouthCarolina in the upper Coastal Plainphysiographic province and occupies78 000 ha (fig. 1). It is located in partsof Aiken, Barnwell, and AllendaleCounties. When the DOE acquired theland in 1950, about 33 percent of it wascropland or pasture, and the remainderwas forested (see Workman andMcLeod 1990 for a detailed descriptionof vegetation communities). In 1952the USDA Forest Service beganmanaging timber on the Site, and 20years later it was designated theNation’s first National EnvironmentalResearch Park (Workman and McLeod1990). The Site has a temperate climatecharacterized by mild winters andlong summers with average airtemperatures of 9

and 27

oC,

respectively (Workman and McLeod1990). The frost-free period lastsroughly 240 days, and the averageannual rainfall is 120 cm. Most rainfalls in March (13.1 cm) and the leastin November (5.9 cm) (Workman andMcLeod 1990).

Using Workman and McLeod (1990)and Imm and McLeod (in press),we identified seven major vegetationtypes on the Site (fig. 2). These includeloblolly-slash pine (Pinus taeda L., P.elliottii Engelm.) (32 461 ha), longleafpine (P. palustris Mill.) (17 008 ha),mixed pine-hardwood (4569 ha),upland hardwood (2738 ha), andbottomland hardwood (16 462 ha)forests; lakes/ponds/marshes (1672ha), and grass-brush (5214 ha).

Pine forests dominate the SavannahRiver Site landscape, and pine canopiesthere tend to be dominated by a singlespecies. Because of their age andmanagement on the Site, longleaf pinestands 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/ RockSpecies Foliage cavity structure mine crevices

Eastern pipistrelle S S S WSSoutheastern myotis S S WSLittle brown bat S S WSSmall-footed myotisa S W SNorthern long-eared myotisa S WS WEvening bat S S SRafinesque’s big-eared bat WS WS WSSilver-haired bat WS W W SEastern red bat WSSeminole bat WSNorthern yellow bat WSHoary bat WSBig brown bat WS WS WSBrazilian 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 Ridge2 - Piedmont3 - Upper Coastal Plain3 - Lower Coastal Plain

turkey oak (Quercus laevis Walt.),scrub post oak (Q. margaretta Sarg.),bluejack oak (Q. incana Bartr.),blackjack oak (Q. marilandicaMuenchh.), blackgum (Nyssa sylvaticaMarsh.), sand hickory [Carya pallida(Ashe) Engl. & Graebn.], mockernuthickory [C. tomentosa (Poir.) Nutt.],sassafras [Sassafras albidum (Nutt.)Nees], and sparkleberry (Vacciniumarboreum Marsh.) (Workman andMcLeod 1990). On more mesic uplandand bottomland sites, loblolly pineforms mixed stands with the hardwoodspecies typical of such sites.

Upland hardwood communities onthe Site are dominated by oaks andhickories and are located on the flats ofthe Aiken Plateau, on stream bluffs,

basal areas than loblolly and slash pineforests. Because vegetation density isa primary factor affecting bat foragingactivity (Owen and others, in press),we treated longleaf forest separately(fig. 2). Both forest types are managedfor timber production on the Sitewithin the constraints of red-cockadedwoodpecker (Picoides borealis Vieillot)habitat management guidelines.

1

See Blake (in press) for a detaileddiscussion of forest managementpractices there. Although the rotationlength for pines there is relatively longfor managed forests in the Southeast

(50 to 120 years), many age classesare found in the region. Clearcutharvesting is used to convert slashpine to species better suited for site-specific conditions, i.e., longleaf,loblolly pine. Young pine plantationsconstitute most early successionalhabitat on the Site. Managers thereuse prescribed fire on a 3- to 5-yearrotation to treat areas of pine forest.

Mixed pine-hardwood forests occupymuch of the Savannah River Site,ranging from the dry and infertilesandhills, which are remnant beachdunes from the Cretaceous shoreline,to moist, rich bottomland sites, whichare subject to periodic inundation.Tree species occurring in sandhillcommunities include longleaf pine,

1 Edwards, J.W.; Smathers, W.M., Jr.; Lemaster,E.T.; Jarvis, W.L. 2000. Savannah River Site red-cockaded woodpecker management plan. 66 p.On file with: USDA Forest Service-SavannahRiver, P.O. Box 700, New Ellenton, SC 29809.

Major Vegetation Types of the Savannah River Site


and along the bordersof bottomland forests(Workman and McLeod1990). Upland foreststypically contain mixtures ofwhite oak (Q. alba L.), post oak(Q. stellata Wangenh.), black oak(Q. velutina Lam.), scarlet oak (Q.coccinea Muenchh.), southern redoak (Q. falcata Michx.), blackjackoak, 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 uplandhardwood communities may containtulip poplar (Liriodendron tulipiferaL.), beech (Fagus grandifolia Ehrh.),red buckeye (Aesculus pavia L.),sourwood [Oxydendrum arboreum(L.) DC.], and witch-hazel(Hamamelis virginiana L.). Uplandhardwood communities are denotedas upland hardwoods on the foresthabitat maps included in each speciesaccount (fig. 2).

Bottomland hardwood forests onthe Site are typical of the mesicmixed-hardwood forests of thesoutheastern Coastal Plain (Workmanand McLeod 1990). Bottomlandhardwood communities occur in thelowlands at elevations between 25and 90 m and commonly are floodedduring high-water periods in latewinter and early spring (Workmanand McLeod 1990). Most hardwoodson the Site were selectively harvestedin the late 1800s and early 1900s.They remained relatively undisturbed

Figure 2—The 449 locations where AnaBat surveys were conducted during summer2001, and the distribution of 7 habitat types across the Savannah River Site.

during the last three-quarters of the20

th century (Workman and McLeod

1990). Common tree species in thebottomland forests include sweetgum,red maple (Acer rubrum L.), swampchestnut oak (Q. michauxii Nutt.),cherrybark oak (Q. falcata var.pagodifolia Ell.), water oak (Q. nigraL.), laurel oak, willow oak (Q. phellosL.), overcup oak (Q. lyrata Walt.),green ash (Fraxinus pennsylvanicaMarsh.), and blackgum. Because thestructure of most swamp forests onthe Site—occurring primarily along

the Savannah River (Whipple andothers 1981)—is similar to thestructure found in bottomlandhardwood forests, we displayedthem as bottomland hardwoodson the forest habitat maps includedwith each species (fig. 2), and weotherwise included them withbottomland hardwood communities.Swamp forests occur at elevationsbetween 25 and 35 m and commonlyare inundated during a portion of theyear (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-growthbald cypress [Taxodium distichum(L.) Rich. var. distichum])-watertupelo (N. aquatica L.) swamp, andthe other half is composed of ridgesand hardwood islands (Jensen andothers 1984). Common tree speciesin swamp forest areas include baldcypress, water tupelo, water ash(F. caroliniana Mill.), green ash,American elm (Ulmus americanaL.), and sycamore (Platanusoccidentalis L.).

Most of the acreage in lakes,ponds, and marshes is in thereservoirs of Par Pond and LLake, which the DOEconstructed for use incooling reactor. Althoughnot extensive, an importanthabitat included in this categoryis Carolina bays. Carolina baysare natural shallow elliptically shapedwetlands whose depths range fromzero to a few meters (Schalles andothers 1989). Although they occurfrom Florida to Virginia, mostCarolina bays are in North andSouth Carolina. Typically they areoriented northwest to southeast.Approximately 500,000 Carolina baysoccur on the Coastal Plain, and about300 are on the Savannah River Site(Kirkman and others 1996). MostCarolina bays are surrounded byupland pine-mixed hardwood forest.Bat activity commonly is concentratedover the wetland habitat that Carolinabays provide, because overstoryvegetation is sparse and aquaticinsects congregate around them(Menzel and others 2003). BecauseCarolina bays are small and occurin patchy distribution across thelandscape, it is hard to map thepatterns of bat activity relative tothem (see figure 3 for the location ofCarolina bays). To simplify the forest-habitat maps included with eachspecies account, we did not identifyCarolina bays, although we didinclude them in the figures thatcompare bat activity levels amongthe habitat types.

Grass-brush habitats include roads,railroad and utility rights-of-way, andlawns around the Site’s facilities areas.The facilities themselves are shownon the vegetation maps. Such grassyareas constitute the only permanentlymaintained early successional habitaton the Site. Typically, the rights-of-wayare dominated by a mix of broomsedge(Andropogon spp.), blackberry (Rubus

spp.), Japanese honeysuckle(Lonicera japonica), andvarious grasses.

Because of their importanceto foraging bats, we delineatedseparately the sampling points locatedwithin canopy gaps (regardless ofvegetation type) and included themon maps displaying the Carolina baysand seven vegetation types.

Major Vegetation Types of the Savannah River Site


The Sitewide Bat Survey

In preparing the species accounts,we relied heavily on a surveyconducted during summer 2001.

In that survey we sampled 449 pointsacross the Site, covering areasrepresentative of most site habitats(fig. 2). We conducted the surveysbetween sunset and 02:30 from lateMay to early August 2001 [see Menzel(2003) for a detailed description of themethods]. At each sampling point, wecensused bat activity for 20 minutesusing an AnaBat II bat detectionsystem (Titley Electronics, Ltd.,Ballina, New South Wales, Australia)(Murray and others 2001). Weidentified calls using both qualitativeand quantitative techniques (Menzel1998, O’Farrell and others 1999) anda call library, which we created byrecording the calls of hand-releasedbats captured in the Southeast. Figures4 and 5 are illustrations generatedusing AnaLook software (4.8i), andthey represent typical search phasecalls of the nine bat species occurringregularly on or around the Site.Menzel (2003) was not able todiscriminate between the calls ofeastern red (Lasiurus borealis) andSeminole (L. seminolus) bats based onthe calls recorded, so both species areincluded in a group termed “easternred/Seminole bat.” We determined theage and vegetation type of the standaround each survey point using 1985Forest Inventory and Analysis dataprovided by the USDA Forest Service.

The following results are condensedfrom Menzel (2003). Trends in batactivity were relatively consistentamong habitat types. The activity level(number of calls per 20-minute survey)of the eastern red/Seminole bat groupwas highest, followed by easternpipistrelles, evening bats, big brownand hoary bats, and southeasternmyotis (in order of decreasing activity)(fig. 6). Because of the highly variablenature of bat activity across the Site, wefound few differences in the levels ofactivity among species within eachhabitat type (table 4). Overall, we Ta

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s th

at o

ccu

r o

n o

r ar

ou

nd

the

Sava

nn

ah R

iver

Sit

e: (A

) eas

tern

red

/Sem

ino

le b

at, (

B) e

ven

ing

bat

, (C

) eas

tern

pip

istr

elle

s, a

nd

(D) s

ou

thea

ster

n m

yoti

s.

9

0123456789

Lake

s and

pon

dsCar

olina

bay

s

Botto

mlan

d ha

rdwoo

ds

Fores

t gap

s

Grass

-bru

sh Upland

har

dwoo

dsPine

-har

dwoo

ds Loblo

lly-s

lash

pines

Long

leaf p

ines

Hab

itat t

ypes

Mean number calls per 20 minute survey

Eas

tern

red

/Sem

inol

e ba

t

Eas

tern

pip

istr

elle

Eve

ning

bat

Big

bro

wn

bat

Hoa

ry b

at

Sou

thea

ster

n m

yotis


Fig

ure

6—

Effe

ct o

f hab

itat

typ

e o

n th

e fl

igh

t act

ivit

y o

f six

bat

sp

ecie

s o

n th

e Sa

van

nah

Riv

er S

ite.

‘


0123456789

101112131415

Lake

/pon

d

Caroli

na b

ays

Botto

mlan

d ha

rdwoo

d

Fores

t gap

s

Grass

/bru

sh

Upland

har

dwoo

d

Pine-h

ardw

ood

Loblo

lly-s

lash

pine

Long

leaf p

ine

Habitat types

Mea

n nu

mbe

r of

cal

ls p

er

20 m

inut

e su

rvey

A

B

Figure 7—(A) AnaBat survey locations where bat activity was detected in surveys conducted during summer 2001and (B) a comparison of bat flight activity among vegetation community types on the Savannah River Site.

11

Tabl

e 5—

Com

paris

on o

f flig

ht a

ctiv

ity (n

umbe

r of c

alls

per

20-

min

ute

surv

ey) a

mon

g si

x ba

t spe

cies

in fi

ve st

and-

age

clas

ses o

n th

e Sa

vann

ah R

iver

Site

durin

g su

mm

er 2

001

Easte

rn re

d/Ea

stern

Sout

heas

tern

Surv

eySe

min

ole b

ats

pipi

stre

lles

Even

ing

bats

Big

brow

n ba

tsHo

ary b

ats

myo

tis

Age c

lass

loca

tions

Mea

nSE

Mea

nSE

Mea

nSE

Mea

nSE

Mea

nSE

Mea

nSE

FP

no.

Clea

rcut

65.

32.

553.

33.

332.

00.

930.

80.

650.

0.0.

20.

175.

300.

001

4–20

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

0.00

121

–40

363.

0A1.

040.

4B0.

220.

4B0.

210.

1B0.

070.

0B0.

031.

0B0.

4922

.27

0.00

141

–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

0.03

2>

60

663.

4A0.

701.

7AB

0.94

1.1B

0.50

0.4B

0.24

1.3B

1.20

0.3B

0.15

1.37

0.26

4

Mea

ns fo

llow

ed b

y the

sam

e let

ter w

ithin

a co

lum

n ar

e not

sign

ifica

ntly

diffe

rent

(P ≤

0.0

5).

detected bat activity on 87.5 percent ofthe lake and pond survey points, 80.0percent of the Carolina bay surveypoints, 66.7 percent of the bottomlandsurvey points, 71.1 percent of theforest gap survey locations, 74.6percent of the grass-brush surveypoints, 63.6 percent of the uplandhardwood points, 57.1 percent of thepine-hardwood points, 48.6 percent ofthe loblolly-slash survey points, and58.6 percent of the longleaf surveylocations. Sitewide, we recorded batactivity at 62.6 percent of all surveylocations (fig. 7). For all speciescombined, activity was highest ingrass-brush and bottomlandhardwoods and over lakes, ponds,and Carolina bays. Levels of activitytended to be low in upland hardwoodand pine-hardwood and on sites withpine-dominated canopies (fig. 7).

In all forest age classes exceptclearcut, bat activity levels alsodiffered among the species (table 5).This trend was consistent among ageclasses other than clearcut. The levelof activity of the eastern red/Seminolebat group was highest, followed byeastern pipistrelles, evening bats,big brown and hoary bats, andsoutheastern myotis (table 5). For allspecies combined, activity was highestin clearcuts, moderate in stands aged 4to 20 years and > 60 years, and lowestin stands aged 21 to 60 years (fig. 8).This trend corresponds with thetypical density of vegetation in standsin these age classes. In clearcuts, batforaging activity is not impeded byforest clutter. The area above newlyregenerating stands (4 to 20 years)provides excellent foraging habitat.Stands aged 21 to 60 years often arestocked so densely that there is littleroom for bats to forage. After standsreach > 60 years, their canopies beginto open—either through natural standattrition or as a result of thinningtreatments—and these openingsprovide foraging areas.



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)

Mea

n nu

mbe

r ca

lls p

er 2

0 m

inut

e su

rvey

0

1

2

3

4

5

6

7

8

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

Age class (year)

Mea

n nu

mbe

r ca

lls p

er 2

0 m

inut

e su

rvey

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 twospecies did not correspond to theactivity pattern of all species combined(fig. 9). Hoary bat activity was highestin stands aged > 60 years, and nohoary bat calls were detected inclearcuts (fig. 9). Southeastern myotisactivity was highest in stands 21 to 40years old and lowest in the youngestand oldest stands.

Accounts of Species

Species accounts are organizedinto two major sections, “Batsof the Savannah River Site” and

“Bats of South Carolina in AreasNeighboring the Savannah RiverSite.” Twelve bat species have beendocumented on the Site, but not alloccur there regularly. To determinethose species that do, we examined therange of each of the 14 bat speciesfound in South Carolina (Menzel andothers 2003), reviewed museumrecords for specimens captured onthe Site itself, and reviewed capturerecords maintained for bat surveystaken there. In summarizing previouscapture and collection records, wereviewed museum records listedin Menzel and others (2003) forspecimens captured on the Site;recorded unpublished survey recordsfrom 1999 to 2001 for one of theauthors;

2 recorded capture records

reported by Carter (1998), Menzel(1998), and Childs and Buchler (citedin Cothran and others 1991); reviewedrecords maintained by public healthpersonnel in South Carolina, asreported by DiSalvo and others(2002); and reviewed recordsmaintained by the South CarolinaDepartment of Natural Resources.Based on our review, some speciescaptured on the Site probably donot occur there regularly and shouldbe of little concern to Site managers,e.g., the northern yellow bat (L.intermedius) and the little brownbat (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, wetried to synthesize all informationcollected in South Carolina aboutroosting ecology, foraging behavior,diet, home range, and reproduction.We used information from publishedstudies, faunal surveys of specificregions within the State (Cothran andothers 1991, Golley 1966, Penney 1950,Sanders 1978), unpublished master’stheses (Carter 1998, Menzel 1998),and unpublished museum records.Because aspects of the natural historyof some bat species found on the Sitehave not been studied anywhere inSouth Carolina, we supplementedinformation in each species accountwith information collected from otherregions. Where information collectedfrom other regions was used, we notedit in the individual species accounts.

In each species account, weincluded the following subsections—Morphology and Distribution,Roosting, Foraging and Home Range,Effect of Stand Age and Habitat Typeon Flight Activity, and Reproduction.Morphology and Distributiondescribes the species’ general physicalcharacteristics, subspecies that occuron the Site, and species distribution.That section includes a picture of thespecies and a table providing itsaverage body measurements. Dataprovided in the body measurementtable were collected from specimenscaptured or collected in the Southeast(primarily Georgia). If the averagebody measurements differedsignificantly (t-test, alpha < 0.05)between males and females, weprovided a separate table for each sex.Each Morphology and Distributionsection references a map illustratingdistribution of the species in SouthCarolina, adapted from Menzel andothers (2003).

The Roosting section includesinformation about a species’ summerand winter roosting habits.

Information about the general roostinghabits of each species also is provided.In addition, specific informationconcerning roosting habits in SouthCarolina or on the Site is provided forseven of the nine bat species thatregularly occur on-site.

The Foraging and Home Rangesection includes information aboutemergence time, foraging habitats,specific foraging characteristics, e.g.,foraging height, whether the specieshas ever been documented foraging ingroups, and diet. When data wereavailable, we included informationabout selective foraging. Also whereavailable, we included informationabout the spatial attributes and size ofthe species’ home range. We definedhome range as the spatial extent anindividual traveled during 3 to 15nights. All reported home ranges werebased on > 25 telemetry points andmost were based on > 30 points.

In each species account, we discusshabitat types on the Site in which aspecies’ foraging activity wasconcentrated and the habitat types itseemed to use less. This information isbased largely on the sitewide surveysconducted during summer 2001. Weused survey data to prepare mapsshowing the general distribution ofeach species’ foraging activity acrossthe Site. We also prepared figures foreach species or species group, i.e.,eastern red/Seminole bat, that showthe average level of call activity in eachvegetation type.

We conclude each account witha brief discussion of the species’reproductive habits. Because littlework has been done with regardto bat reproductive habits in SouthCarolina, most of this informationcame from studies conducted inother regions of the United States.Information included in this sectionincludes the time of year matingoccurs, whether delayed fertilizationoccurs, when the young are born, andthe 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


Bats of theSavannah River Site

FAMILY VESPERTILIONIDAE

Vespertilionidae is the largestfamily of bats, consisting of42 genera and 355 species

(Nowak 1994). Family distributionis cosmopolitan, and members arefound on every continent exceptAntarctica. Its members are foundin habitats ranging from desert totropical forest. Because members ofthis group lack noseleafs and havesimple, unmodified lips and nostrils,they are commonly called the plain-faced bats. The tragus usually is welldeveloped, and the tail is not free fromthe uropatagium.

Nine genera and thirty-one speciesof vespertilionids occur in the UnitedStates. Of these, 8 genera and 13species occur in South Carolina.All nine bat species that commonlyoccur on the Savannah River Site aremembers of the family Vespertilionidae.

Eastern pipistrelle(Pipistrellus subflavus)

Morphology and DistributionThe eastern pipistrelle is the second

smallest bat found in South Carolinaand the smallest bat on the Site(fig. 10). One of four recognizedsubspecies, P. s. subflavus, occurs inthe State (Fujita and Kunz 1984). Massranges from 3.3 to 8.0 g, and averagetotal length is 85.1 mm (Whitaker andHamilton 1998). See table 6 for averagebody measurements of individualscollected in the Southeast.

The eastern pipistrelle is commonthroughout much of the eastern halfof North and Central America. In theUnited States, its range extends westfrom Maine to Minnesota and souththrough much of east Texas (Fujitaand Kunz 1984). Thirty-nine easternpipistrelles from South Carolina arepreserved in collections, and 23 havebeen live-captured in the State (Menzel

and others 2003). The easternpipistrelle occurs in all of SouthCarolina’s physiographic provincesand is common throughout the State(fig. 11) (Menzel and others 2003).

Eastern pipistrelles were capturedin late June and late August duringsurvey work conducted by Childs andBuchler in 1979 (Anon. 1980, Cothranand others 1991). Between 1996 and2001, we captured one female and five

male eastern pipistrelles on the Site(table 7). Based on the regionaldistribution of this species, it is likelythat while it is not abundant on theSite, the eastern pipistrelle is acommon resident there.

RoostingEastern pipistrelle hibernacula have

been found in culverts (Moore 1949),storm sewers (Goehring 1954), tunnels(Mohr 1942), caves (Davis 1966, Hahn

Figure 10—Eastern pipistrelle.

Phot

o co

urte

sy o

f Joh

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acG

rego

r

15

Figure 11—Distribution of the eastern pipistrelle in South Carolina (Menzeland others 2003). Symbols denote counties for which records exist.

Eastern pipistrelle(Pipistrellus subflavus)

Table 6—Body measurements of the eastern pipistrelle bat in the SoutheasternUnited States

Measurement Numbera Mean SD CV Range

- - - - - - - - - - - - - - - - - - mm - - - - - - - - - - - - - - - - - -Female

Total 15 86.3 5.96 6.90 77.0 – 99.0Tail 15 39.0 3.16 8.11 34.0 – 45.0Foot 15 7.8 1.21 15.48 6.0 – 10.0Ear 15 12.8 1.56 12.16 10.0 – 14.5Tragus 14 4.7 1.37 28.98 3.5 – 7.5Forearm 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


- - - - - - - - - - - - - - - - - - - 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, Swansonand Evans 1936), and mines (Menzeland others 1997, Sealander and Heidt1990, Whitaker and Rissler 1992). Nostudies have quantitatively examinedits winter roosting habits in SouthCarolina, although Golley (1966)collected a male eastern pipistrelleroosting under a house in BerkeleyCounty, SC, during the winter. Golley(1966) also found a large colony ofeastern pipistrelles roosting in cavesnear Parler in Orangeburg County,SC, during the winter and spring.The winter roosting habits of thisspecies on the Savannah River Siteare unknown.

The eastern pipistrelle’s summerroosting habits also are poorly known.Maternity roosts usually are found inhuman-made structures such as barns(Lane 1946, Poole 1938) or houses(Allen 1921). Menzel and others (1996)captured a female and her nonvolantyoung in a pitfall trap set near a basalcavity in a large sweetgum, whichsuggests that basal cavities may serveas maternity roosts for the species. Afew individuals also have been foundroosting in tree canopies (Findley1954) and Spanish moss (Tillandsiausneoides) during the summer(Jennings 1958, Menzel and others1999). In summer 1997, Carter andothers (1999b) located an easternpipistrelle roosting in swamp chestnutoak, sweetgum, and laurel oak in abottomland hardwood stand onthe Site.


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 0M 5 0

Southeastern myotis (n = 2) F 0 0M 1 0

Little brown bat (n = 1) F 1 0M 0 0

Evening bat (n = 67) F 27 13M 21 6

Rafinesque’s big-eared bat (n = 9) F 7 0M 1 0

Eastern red bat (n = 56) F 30 11M 8 5

Seminole bat (n = 22) F 6 6M 7 3

Hoary bat (n = 3) F 2 0M 0 0

Big brown bat (n = 6) F 2 1M 3 0

Total (n = 173) F 70 31M 52 14

n = total number captured (the difference between n and sum of age and sex data isbecause age and sex were not recorded for all individuals captured);F = female; M = male.

Foraging and Home RangeEastern pipistrelles commonly

forage over waterways and alongfield edges (Fujita and Kunz 1984,Whitaker and Hamilton 1998). Harper(1927) found them foraging over fieldsand cypress bays and in pine barrensof the Okefenokee Swamp. On theSavannah River Site, Carter and others(1999b) documented foraging amongbottomland hardwoods and in pinestands. Their diet includes insects inthe orders Coleoptera, Homoptera,Diptera, Hymenoptera, and Lepidoptera(Ross 1961, Sherman 1939, Whitaker1972). They also forage ontrichopterans and hemipterans(Carter and others 1999b).

Little is known about this species’home range, although Krishon andothers (1997) documented the homerange of one eastern pipistrelle onSapelo Island, GA. It was 389.2 ha andconcentrated in high marsh (47percent), oak (24 percent), and slash-loblolly pine (17 percent) habitats. Theaverage distance from the roost area toa foraging location was 1137 m. Carterand others (1999b) documented aneastern pipistrelle’s home range onthe Site at 395.5 ha, concentrated inbottomland hardwood and uplandpine stands.

Effect of Habitat Type and StandAge on Flight Activity

We recorded eastern pipistrelles at37.5 percent of the lake and pondsurvey points, 28.6 percent of theCarolina bay survey points, 18.2percent of the bottomland hardwoodsurvey points, 18.4 percent of theforest gap survey locations, 23.7percent of the grass-brush surveypoints, 9.1 percent of the uplandhardwood points, 21.4 percent of thepine-hardwood points, 12.1 percent ofthe loblolly-slash survey points, and7.1 percent of the longleaf surveylocations. Overall, we recorded theeastern pipistrelle’s call at 16.7 percentof the survey locations (fig. 12).Although variable, its activity wasgreatest around lakes and ponds,bottomland hardwood forests, andgrass-brush (fig. 12). In bottomlandhardwoods, we recorded a callapproximately every 10 minutes. Thefigures comparing bat activity amongvegetation community types arearranged with hydric communities onthe left, mesic communities in thecenter, and xeric communities on theright. Eastern pipistrelle activity wasconcentrated in hydric and mesiccommunities (table 4).

Eastern pipistrelle activity alsodiffered among stands of differentages (fig. 9). We recorded most inclearcuts (fig. 9) or areas devoid ofvegetation, e.g., roads, open waterhabitats (fig .12). We recordedmoderate levels of activity in standsaged 4 to 20 years and > 60 years (fig.9). The species’ activity was low instands 21 to 60 years old (table 5).

ReproductionEastern pipistrelles mate in autumn

and spring (LaVal and LaVal 1980) andparturition occurs in May and June(Jennings 1958, LaVal and LaVal 1980).The sexes segregate into bachelor andmaternity colonies in the spring. Noinformation is available concerning theparturition times for this species inSouth Carolina, but on the SavannahRiver Site it probably occurs in Mayand June.

17

0.0

0.5

1.0

1.5

2.0

2.5

3.0

3.5

4.0

4.5

Lake

/pon

d

Caroli

na b

ays

Botto

mlan

d ha

rdwoo

d

Fores

t gap

s

Grass

/bru

sh

Upland

har

dwoo

d

Pine-h

ardw

ood

Loblo

lly-s

lash

pine

Long

leaf p

ine

Habitat types

Mea

n nu

mbe

r of

cal

ls p

er20

min

ute

surv

eyA

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.



Table 8—Body measurements of the southeastern myotis bat in theSoutheastern United States


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


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

Mass 12 5.7 0.79 13.91 4.3 – 7.0

a Number of individuals measured.

Southeastern myotis(Myotis austroriparius)

Morphology and DistributionThe southeastern myotis is a small

brown bat with wooly pelage anda calcar that lacks a keel (fig. 13)(Jones and Manning 1989). Of thethree subspecies of southeasternmyotis now recognized, only M. a.austroriparius occurs in South Carolina(Hoffmeister 1989). The mass of thesoutheastern myotis ranges from 5 to12 g, and total length averages 91.5mm (Whitaker and Hamilton 1998).See table 8 for average bodymeasurements of individuals collectedin the Southeast.

The southeastern myotis’ rangeextends from Louisiana and east Texasup the Mississippi Alluvial Valley intosouthern Indiana and Illinois, and eastalong the coast of Georgia and theCarolinas (Jones and Manning 1989).Four southeastern myotis from SouthCarolina are in museum collections,

Figure 13—Southeastern myotis.

and 18 have been live-captured.Distribution in South Carolina islimited to the upper and lower CoastalPlain (fig. 14). Because of its raritythere, the State has listed the speciesas threatened (South CarolinaDepartment of Natural Resources2001) (table 9).

Only two southeastern myotis havebeen captured on the Site, one maleand one sex undetermined, captured in1997 and 2000, respectively (table 7).None was captured during the 1979surveys. The Site is on the northernedge of the species’ range, andalthough the two documented records

Phot

o co

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rego

r.

19

Southeastern myotis(Myotis austroriparius)

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

South Carolina

Species State Protectionranking status Abundance

Eastern pipistrelle CommonSoutheastern myotis S1 ST RareLittle brown myotis S3 SC UncommonSmall-footed myotisb S1 ST RareNorthern long-eared myotisb S4 SC CommonEvening bat CommonRafinesque’s big-eared bat S2 SE RareSilver-haired bat UncommonEastern red bat CommonSeminole bat CommonNorthern yellow bat SU SC RareHoary bat SU SC UncommonBig brown bat CommonBrazilian 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).


Figure 14—Distribution of the southeastern myotis in South Carolina (Menzeland others 2003). Symbols denote counties for which records exist. Typical

summer distribution is south of the range line.

probably are not extralimital, it appearsto be rare on the Site.

RoostingSoutheastern myotis roost in caves

(Mumford and Whitaker 1982, Rice1957), mines (Sealander 1979), trees(Lowery 1974, Sealander 1979), andbuildings (Lowery 1974, Mumford andWhitaker 1982, Sealander 1979). Davisand Rippy (1968) banded a colony of300 individuals roosting in a fertilizerplant in Georgia. Only Clark andothers (1998) have investigated theroosting habits of southeastern myotisin South Carolina. Roosts were foundin cavities of live tupelo gum trees in aclosed-canopy forest on the FrancisBeidler Forest (Clark and others1998). Based on this information andthe species’ roosting habits in otherareas of its range, we believe itprobably roosts in tree cavities inbottomland hardwood and swampforests on the Site.

Foraging and Home RangeSoutheastern myotis typically forage

near water (Barbour and Davis 1969).In the winter they prey on arthropodsin the orders Coleoptera andLepidoptera and in the family Culicidae(Zinn 1977). During the summermonths they consume primarilyColeoptera and Lepidoptera (Zinn1977). Clark and others (1998)surveyed too few foraging locations todetermine home-range size, but allforaging points were in swamp forests.We know little about the size of thespecies’ home range or habitat use onthe Site. However, based on the knownhome range of other myotids withsimilar morphological characteristics,we surmise the southeastern myotis’home range is between 100 and 500 ha.Data collected during the 2001 AnaBatsurveys suggest that foraging activityon the Site is concentrated in swampforest bordering the Savannah River(fig. 15).

Effect of Habitat Type andStand Age on Flight Activity

Southeastern myotis were recordedat 12.5 percent of the lake and pondsurvey points, 22.9 percent of the


0.0

0.5

1.0

1.5

Lake

/pon

d

Caroli

na b

ays

Botto

mlan

d ha

rdwoo

d

Fores

t gap

s

Grass

/bru

sh

Upland

har

dwoo

d

Pine-h

ardw

ood

Loblo

lly-s

lash

pine

Long

leaf p

ine

Habitat types

Mea

n nu

mbe

r of

cal

ls p

er20

min

ute

surv

ey

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

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r

Carolina bay survey points, 13.6percent of the bottomland hardwoodsurvey points, 26.3 percent of theforest gap survey locations, 5.1 percentof the grass-brush survey points, 9.1percent of the upland hardwood points,7.1 percent of the pine-hardwoodpoints, 7.1 percent of the loblolly-slashsurvey points, and 2.9 percent of thelongleaf survey locations. Overall, werecorded this species at 10.2 percentof the survey locations (fig. 15). Its on-site foraging activity was low (table 4).The mean number of calls recordedwas less than one per 20-minutesurvey in all habitat types (fig. 15).Activity was highest in Carolina bays,bottomland hardwood forests, andforest gaps; and the survey pointswhere southeastern myotis weredetected were concentrated nearswamp forests along the SavannahRiver (fig. 15).

Southeastern myotis’ foragingactivity varied among stands of

different ages (fig. 9). Unlike theeastern pipistrelle’s, most of thisspecies’ activity was in stands aged 21to 40 years (table 5).

ReproductionSoutheastern myotis mate in

autumn; fertilization and parturitionoccur in spring (Lowery 1974). InFlorida, Rice (1957) found that thespecies entered maternity roostsaround mid-March. Most maternityroosts were in caves and containedbetween 2,000 and 90,000 individuals ataverage densities of 1,600 bats per m

2.

Young were born from late April tomid-May. The southeastern myotisis the only myotid in South Carolinathat typically gives birth to twins(Barbour and Davis 1969). Althoughno maternity colonies have beenstudied in South Carolina, parturitionprobably occurs in May and early Juneon the Savannah River Site.

Evening bat(Nyticeius humeralis)

Morphology and DistributionThe evening bat is a small brown

bat that occurs throughout theSoutheastern United States (fig. 16).Of the three recognized subspecies,only one, N. h. humeralis, occursin South Carolina. Weight ranges from5 to 14 g, and average total length is92.7 mm (Whitaker and Hamilton1998). See table 10 for average bodymeasurements of individuals collectedin the Southeastern United States.

The evening bat’s range extendsnorth from the Texas-Mexico borderto Nebraska, and east throughPennsylvania (Watkins 1972). Museumcollections include 172 specimens fromSouth Carolina; 93 have been live-captured in the State. The evening batoccurs in all physiographic provincesof South Carolina and is commonstatewide (fig. 17) (Watkins 1972).


Figure 16—Evening bat.


Evening bat(Nycticeius humeralis)

Table 10—Body measurements of the evening bat in the SoutheasternUnited States


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

Total 20 86.8 6.28 7.23 77.0 –102.0Tail 20 33.8 4.07 12.04 26.0 – 39.0Foot 20 7.9 1.34 16.97 6.0 – 11.0Ear 19 11.9 2.02 16.96 6.5 – 14.5Tragus 20 3.9 1.38 35.14 2.5 – 7.0Forearm 18 36.2 1.81 4.99 33.0 – 39.0

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

Mass 18 8.0 2.04 25.57 4.3 – 12.4


Figure 17—Distribution of the evening bat in South Carolina (Menzel and others2003). Symbols denote counties for which records exist.

23

Evening bats were captured duringthe 1979 survey on the Site (Anon.1980, Cothran and others 1991). In on-site surveys conducted from 1996to 2001, it was the most commonlycaptured species; we captured 27 adultfemales, 21 adult males, 13 juvenilefemales, and 6 juvenile males (table 7).

RoostingNo studies have quantitatively

examined winter roosting habits ofevening bats in South Carolina, andlittle is known about their winterroosting habits anywhere. There are,however, anecdotal accounts ofevening bats roosting in attics inLegareville, Charleston County, SC,during the winter.

Menzel and others (2000b) found 61evening bat summer roost trees on theSite. The most common tree specieswas longleaf pine, but conifer snags inbeaver ponds also were common(Menzel and others 2000b). Eveningbats roosted either in cavities or underexfoliating bark. Roosts were in areaswith fewer overstory and understorytrees, greater average canopy height,lower understory richness anddiversity, lower overstory richness,less-dense canopies, and greater snagabundance than in random plots(Menzel and others 2000b). Eveningbat maternity colonies used roosts inmature longleaf pine in stands wherethe height of the overstory, density ofthe canopy, and proportion of the basalarea composed of conifers weregreater than in areas surroundingroost trees used by solitary eveningbats (Menzel and others 2001a).Summer roosts have been foundelsewhere in Spanish moss (Jennings1958), under exfoliating bark (Bailey1933, Barbour and Davis 1969,Chapman and Chapman 1990), and intree cavities (Barbour and Davis 1969,Harper 1927). There is one record ofthis species roosting in caves (Easterla1965). Maternity roosts have been

found in buildings (Cope and others1961, Watkins and Shump 1981) andtree cavities (Menzel and others2001a). In the upper and lower CoastalPlain of Georgia, the species oftenshares maternity roosts in buildingswith T. brasiliensis.

3

Foraging and Home RangeEvening bats begin foraging in late

twilight (Lowery 1974). At firstemergence, they forage 13 to 25 mabove the ground, but later in the nightthey forage lower (Harper 1927,Lowery 1974). Harper (1927) reportedthat in the Okefenokee Swamp area ofGeorgia, evening bats foraged in pinebarrens and over fields. Krishon andothers (1997) found that 76 percent ofthe bats’ foraging area on SapeloIsland, GA, was over slash-loblolly pinehabitat. Evening bats feed on insects inthe orders Hemiptera, Hymenoptera,Coleoptera, and Lepidoptera(Mumford and Whitaker 1982, Ross1961). Carter (1998) found that on theSite they fed primarily on Coleopteraand Hymenoptera during early andmidsummer. In late summer, thespecies fed on a wider variety of insecttaxa, including Hemiptera, Homoptera,Coleoptera, and Hymenoptera(in order of decreasing volume)(Carter 1998).

The home range of one juvenile andfive adult evening bats tracked on theSite ranged from 38.7 to 761.0 ha andaveraged 285.3 ha (Carter 1998).Habitat types contained within thehome ranges included pine forests (59percent) and bottomland hardwoods(37 percent) (Carter 1998). Habitattypes were used in the sameproportion as they were available.The 761.0-ha home range of thejuvenile was much larger than that ofthe other five. On Sapelo Island, GA,the species’ home range was within a15.1-ha slash-loblolly pine stand(Krishon and others 1997).



We recorded evening bats at 31.3percent of the lake and pond surveypoints, 42.9 percent of the Carolina baysurvey points, 31.8 percent of thebottomland hardwood survey points,31.6 percent of the forest gap surveylocations, 40.7 percent of the grass-brush survey points, 27.3 percent ofthe upland hardwood points, 14.3percent of the pine-hardwood points,10.7 percent of the loblolly-slashsurvey points, and 21.4 percent of thelongleaf survey locations. Overall, theywere present on 24.9 percent of allsurvey locations (fig. 18). Most batactivity was within grassy areas,among bottomland hardwoods, andover Carolina bays (fig. 18). Whereaseastern pipistrelle activity was highestover lakes and ponds, evening batactivity was highest over Carolinabays. Unlike the southeastern myotis,evening bat activity was widespreadthroughout the Site.

Stand age also influenced evening batactivity (table 5). As with the easternpipistrelle, activity was highest inclearcuts 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).

ReproductionIn the Southeastern United States,

evening bats give birth in May andJune. In the Northeast, parturition mayoccur in July (Harper 1927, Watkins1972). Evening bats arrive at nurseryroosts around the second week of April(Golley 1966) in South Carolina. InMay, Golley (1966) captured severalpregnant females in the State. Mostbats leave their nursery colonies bylate August (Baker 1965). Young batsbecome volant about 20 days afterbirth (Schmidly 1991). Menzel andothers (2000b) found many eveningbat maternity colonies and capturedmany juvenile evening bats on theSavannah River Site.

3 Personal communication. 1999. Jim Ozier,Senior Wildlife Biologist, Georgia Department ofNatural Resources, Wildlife Resources DivisionNongame-Endangered Wildlife Program, 116Rum Creek Drive, Forsythe, GA 31029-6518.


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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.

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Table 11—Body measurements of Rafinesque’s big–eared bat in theSoutheastern United States


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


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

Mass 11 8.6 1.05 12.15 7.5 – 11.1


Rafinesque’s big-eared bat(Corynorhinus rafinesquii)

Morphology and DistributionRafinesque’s big-eared bat is small,

and its most obvious attribute is itsextremely large ears (fig. 19). Bothsubspecies of Rafinesque’s big-earedbat (C. rafinesquii rafinesquii andC. r. macrotis) occur in South Carolina(Jones 1977). Weight ranges from7 to 10 g, and average total length is99.5 mm (Whitaker and Hamilton1998). See table 11 for average bodymeasurements of individuals collectedin the Southeastern United States.

Rafinesque’s big-eared bat is foundthroughout the Southeast. Its rangeextends south from southern Virginia,along the Atlantic Coast into Florida,and west into Oklahoma and eastTexas (Jones 1977). Museumcollections include 65 specimensfrom South Carolina; 30 individuals

have been live-captured in the State.Its distribution in the South extendsthrough the upper and lower CoastalPlain and the Blue Ridge (fig. 20). It islisted as endangered in South Carolina(table 9) (South Carolina Departmentof Natural Resources 2001).

Two Rafinesque’s big-eared batscollected on the Site are reposited inthe University of Georgia Museum ofNatural History (Cothran and others1991). Between 1996 and 2000, wecaptured seven adult females andone adult male (table 7).

Figure 19—Rafinesque’s big-eared bat.



Rafinesque's big-eared bat(Corynorhinus rafinesquii)

(Menzel and others 2001b). Theyselectively foraged there in sapling-stage pine stands (Menzel and others2001b). The results of this study standin stark contrast to the results of astudy conducted by Clark and others(1998) on the Francis Beidler Forestin central South Carolina, where thehome range of three individuals wasbetween 64 and 89 ha, primarily inswamp forests. On-site, this batprobably forages in a wide variety ofhabitats. However, foraging activitymay be concentrated in maturebottomland hardwood and swampforests, brushy communities, and 3-to 5-year-old pine plantations.

Fecal pellet analyses in SouthCarolina indicated the primary taxaconsumed by Rafinesque’s big-earedbats was Lepidoptera (Menzel andothers 2002b). Ellis (1993) collectedfecal pellets from the roost of amaternity colony in North Carolinaand found that the diet was exclusivelycomposed of Lepidopterans (67percent) and Dipterans (33 percent).Most of the flies eaten (94 percent)were tabanids (family Tabanidae).


Because of the low intensity of theecholocation calls produced, activitypatterns of the Rafinesque’s big-earedbat cannot be monitored reliably withbat detectors. We did not attempt todetermine the effect of habitat typeor stand age on the flight activity ofthis species.

ReproductionRafinesque’s big-eared bats seem to

mate in autumn (Hoffmeister andGoodpaster 1963). Animals withswollen testes have been found inAugust (Hall 1963), and reproductivelyactive males were captured in areasaround the Site in September (Menzeland others 2001b). Females entermaternity roosts in May. Each of twomaternity colonies found afterparturition by Hall (1963) containedabout 50 individuals. The species’typical litter size is one. Young areborn 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.

RoostingThe Rafinesque’s big-eared bat’s

winter roosting habits are not wellknown. It has been found roosting inabandoned buildings throughout theyear (Clark 1990). Winter roosts alsohave been found near cave entrances(Barbour and Davis 1969).

During the summer, Rafinesque’sbig-eared bats roost in abandonedbuildings, hollow trees, and under bark(Barbour and Davis 1969, Hall 1963,Jones 1977, Lowery 1974, Menzel andothers 2001b). Typically, summerroosts are located in the twilight zoneof caves and mines or in dimly litsections of buildings (Whitaker andHamilton 1998). Menzel and others(2001b) found male big-eared batsroosting in abandoned houses in AikenCounty, SC, approximately 8 kmnorthwest of the Savannah River Site.We have found some of them roostingin an abandoned barn on the Site. Theyprobably roost under bridges, in treecavities, and in abandoned structuresnear bottomland and swamp forestson the Site. Clark (1990) describedroosting sites in North Carolina as“predominately frame, one-story

[abandoned] homes with plaster wallsand large attics.” Lance and others(2001) found them roosting under32 girder-type bridges in Louisiana.Bridges probably are an importantcomponent of the species’ roostinghabitat in South Carolina. Futuresurveys should determine whichbridges on the Site serve as roosts forthis species. Lance and others (2001)found big-eared bats roosting in hollowblackgum trees. Based on unpublishedmuseum records, several hundredwere reported roosting in a house onBelle Isle in Georgetown County, SC.Clark and others (1998) found someroosting in tree cavities of water tupeloin the Francis Beidler Forest in SouthCarolina. Most roosts were withinbasal cavities. They also found a1-night roost in a dead bald cypress.

Foraging and Home RangeWe know little about the foraging

habits of Rafinesque’s big-eared bats.They do not begin to forage until afterdark and are agile fliers (Schmidly1991). In a study conducted in AikenCounty, SC, the home-range size ofmale big-eared bats ranged from 23.9to 260.5 ha and averaged 93.2 ha

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In South Carolina, maternity colonieshave been found in abandonedbuildings. Parturition probablyoccurs in May and June on theSavannah River Site.

Silver-haired bat(Lasionycteris noctivagans)

Morphology and DistributionThe silver-haired bat is a small bat

with silvery-white frosting, and itoccurs throughout southern Canadaand most of the United States (fig. 21).Weight ranges from 7 to 16 g, andaverage total length is 99.7 mm(Whitaker and Hamilton 1998).See table 12 for average bodymeasurements of individuals collectedin the Southeastern United States.

The silver-haired bat reaches thesouthern limit of its range in theSoutheast (Kunz 1982). Museumcollections include 26 silver-hairedbats from South Carolina. The speciesis distributed statewide (fig. 22),

and individuals have been capturedin all four of South Carolina’sphysiographic provinces. However,its distribution in the State probablydiffers among seasons, and many ofthe individuals may have beencaptured while migrating.

Although one individual collected onthe Site is reposited in the Universityof Georgia Museum of Natural History,none was captured during the 1979 or1996 to 2001 surveys. The species ismigratory (Barclay 1984a, Kunz 1982).Silver-haired bats probably are rareon the Site between late autumn andearly spring and may not occur therein the summer.

RoostingRoosting habits of the silver-haired

bat in South Carolina are unknown.In other States, they are known tohibernate in mines (Baker 1965, Layne1958, Pearson 1962), caves (Baker1965, Beer 1956, Turner 1974), rockcrevices (Frum 1953), buildings (Clark1993, Frum 1953), and trees (Cowan

1933, Jackson 1961). Summer roostshave been found in tree cavities andunder bark (Betts 1996, Mattson andothers 1996, Parsons and others 1986,Vonhof 1996). Because it is unlikelythat silver-haired bats are on-siteduring the summer, the limitedknowledge of the species’ summerroosting habits in the Southeast is oflittle concern.

Foraging and Home RangeJones and others (1973) and Kunz

(1973) found that silver-haired batstypically have two peaks of foragingactivity per night—3 and 7 hours aftersunset. Barclay (1984b) found thatthey forage throughout the night,typically in coniferous or mixeddeciduous forests that are nearwater (Jones 1965, 1966; Kunz1973). They fly slowly and erraticallywhen foraging (Hayward and Davis1964, van Zyll de Jong 1985). Fooditems include Lepidopterans,Homopterans, Dipterans,Hemipterans, Hymenopterans,Coleopterans, and Neuropterans

Figure 21—Silver-haired bat.


Silver-haired bat(Lasionycterius noctivagans)

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


- - - - - - - - - - - - - - - mm - - - - - - - - - - - - - -Female


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

Mass 3 10.4 0.85 8.20 9.5 – 11.2

- - - - - - - - - - - - - - - mm - - - - - - - - - - - - - -Male


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

Mass 7 8.6 1.58 18.39 5.2 – 9.9


(Black 1974; Whitaker and others1977, 1981). The species’ foraginghabits in South Carolina are notknown. Because silver-haired batsprobably do not reside on-site duringthe summer, the limited knowledgeabout its foraging behavior in theSoutheast is of little concern.


We did not record any calls of silver-haired bats during the 2001 AnaBatsurvey. Although they may occuron-site during the winter months,it is unlikely that they are summerresidents there.

ReproductionThe reproductive habits of silver-

haired bats are not well known.Similar to other vespertilionid bats,they probably mate in the autumn(van Zyll de Jong 1985). Ovulationoccurs in April and May (Druecker1972). They give birth to one or twoyoung in June or July (Easterla andWatkins 1970, Kunz 1971, Merriam1884). It is unlikely that parturitionoccurs on the Savannah River Site.

Figure 22—Distribution of the silver-haired bat in South Carolina (Menzeland others 2003). Symbols denote counties for which records exist.

29

from 9.5 to 16 g, and its average totallength is 112.3 mm (Whitaker andHamilton 1998). See table 13 foraverage body measurements ofindividuals collected in the Southeast.

The range of the eastern red batextends south from southern Canadainto Argentina and Chile (Shump andShump 1982a). Museum collectionsinclude 108 specimens from SouthCarolina; 145 have been live-capturedin the State. The species is abundant inall four physiographic provinces of

Eastern red bat(Lasiurus borealis)

Morphology and DistributionThe eastern red bat is medium

sized and brick red (males) or brickred with white frosting (females); itoccurs throughout forested regions ofthe Eastern United States (fig. 23)(Shump and Shump 1982a). Of thethree recognized subspecies of easternred bats, only one, L. b. borealis, occursin South Carolina. Its mass ranges

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South Carolina and is commonthroughout the State (fig. 24).

One eastern red bat was captured atRainbow Bay during the 1979 survey(Anon. 1980). Between 1996 and 2001,we captured 30 adult females, 8 adultmales, 11 juvenile females, and 5juvenile males (table 7). Based on ourcapture data, the eastern red bat is thesecond most abundant bat species onthe Site.

RoostingEastern red bats migrate to the

Southern United States to hibernate.Generally, winter roosts include treebranches and leaf clusters (Barbourand Davis 1969). The species is welladapted for winter survival outside ofcaves and may become active on warmwinter nights (Barbour and Davis1969). Winter roosting habits in SouthCarolina have not been determined. Aswith the other species, we know littleof the relative abundance of easternred bats on the Savannah River Siteduring the winter or of its winterroosting habits.

In the summer, eastern red batsmost commonly are found roosting onsmall branches and leaf petioles in thecrowns of deciduous trees (Barbourand Davis 1969), but they also havebeen found in woodpecker cavities(Fassler 1975) and caves (Myers1960). Menzel and others (1998) found64 eastern red bat roost trees on theSavannah River Site. Roosts typicallywere in hardwood trees larger indiameter at breast height (d.b.h.) andtaller than surrounding trees. Theroost trees were in areas with greateroverstory tree height and basal area,denser overstory canopies, and greateroverstory and understory diversitythan random plots (Menzel and others2000b). Twenty tree species were usedas roosts on the Site, but sweetgum andred maple were the most common(Menzel and others 2000b). Theeastern red bat moves to new roosttrees often, spending an average of 1.2nights at each (Menzel and others1998). Each individual we trackedselected roost trees from a small areaof their foraging range (average roostselection area = 2.6 ha).


Figure 23—Eastern red bat.


Figure 24—Distribution of the eastern red bat in South Carolina (Menzel andothers 2003). Symbols denote counties for which records exist.

Eastern red bat(Lasiurus borealis)

Table 13—Body measurements of the eastern red bat in theSoutheastern United States


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


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

Mass 27 10.0 1.52 15.15 8.0 – 13.0


Foraging and Home RangeEastern red bats typically begin

foraging 1 to 2 hours after sunset,although some individuals maybegin earlier and continue foragingthroughout the night (Kunz 1973).There is some evidence suggestingthey exhibit bimodal foraging activitypatterns (Carter and others 1999a).The species forages at temperaturesas low as 7

oC (LaVal and LaVal 1979).

In South Carolina, eastern red batsfeed mostly on Coleoptera andHemiptera during early summer(Carter 1998), but in mid- and latesummer they feed on a more diverseassemblage of taxa, includingColeoptera, Hemidoptera, Lepidoptera,Homoptera, and Hymenoptera(Carter 1998). Elsewhere, remains ofLepidoptera, Homoptera, Coleoptera,Hymenoptera, and Diptera have beenfound in the stomachs of eastern redbats (Mumford 1973, Ross 1967,Whitaker 1972).

The home range of five eastern redbats tracked on-site ranged from 125.8to 878.5 ha and averaged 453.2 ha(Carter 1998). Habitat types withinits home range included bottomlandhardwoods (55 percent), pine stands(40 percent), and upland hardwoods(5 percent) (Carter 1998). This speciesdid not appear to forage selectively inspecific habitat types; they used eachin proportion to its availability.


Because it is difficult to differentiatecalls of eastern red and Seminole bats,we included both calls in a singlecategory. Although data collected inprevious studies suggest eastern redbats are more likely to forage inbottomland forests than Seminole bats(Menzel 1998), pooling the calls madeit impossible to distinguish foraginghabitat use of the two. We recordedcalls at 56.3 percent of the lake andpond survey points, 71.4 percent ofthe Carolina bay survey points, 45.5percent of the bottomland hardwoodsurvey points, 68.4 percent of theforest gap survey locations, 62.7percent of the grass-brush surveypoints, 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.

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hardwood points, 50.0 percent of thepine-hardwood points, 36.4 percent ofthe loblolly-slash survey points, and47.1 percent of the longleaf surveylocations. Overall, the species groupwas present on 49.7 percent of thesurvey locations (fig. 25). The group’swide distribution across habitat typesand their high occurrence rateprobably resulted from our inclusionof both species. The effect of poolingwould have an even greater effect onthese metrics if they commonlyforaged in different habitats. IfSeminole bats usually roost and foragein upland habitats and eastern red batsin bottomland forests, such poolingwould result in a group common toboth upland and bottomlandcommunities. Also, because theeastern red and Seminole bats arethe second and third most commonspecies on the Site, respectively,it is not surprising that this groupwas detected on half of all 449survey locations.

In addition to being commonlydetected, the group’s activity levelswere relatively high in all habitat types(fig. 25). Most activity was detectedover Carolina bays and grass-brushhabitat (table 4).

Eastern red and Seminole batactivity also seemed to be less affectedby stand age than that of other species(fig. 9). Although activity was greatestwithin clearcuts, it was relativelyconstant among stands aged 4 years orolder (table 5).

ReproductionEastern red bats undergo delayed

fertilization. Mating occurs in Augustand September, and fertilization inspring (Glass 1966, Whitaker andHamilton 1998). Copulation sometimesoccurs in flight (Stuewer 1948).Estimated gestation is between 80 and90 days (Jackson 1961). Its young areborn in late May and June (Lowery1974), and lactation lasts about 38 days(Kunz 1971). The average embryocount is 3.2 (Shump and Shump1982a). Members of the genusLasiurus are the only bats thatregularly give birth to more than two

progeny (Nowak 1994). Although noone has studied the reproductivehabits of eastern red bats in SouthCarolina, we have captured juvenileson the Site, and it is likely that theycommonly raise their young on-site.

Seminole bat(Lasiurus seminolus)

Morphology and DistributionThe Seminole bat is a medium sized,

mahogany colored bat commonthroughout the Deep South (fig. 26).

The Seminole bat is monotypic. Itsmass ranges from 9 to 14 g, andits average total length is 111 mm(Whitaker and Hamilton 1998).See table 14 for average bodymeasurements of individuals collectedin the Southeast.

Although a few “wandering”individuals have been captured in NewYork (Layne 1955) and Pennsylvania(Poole 1949), this species typically isfound south from the southeastern tipof Virginia to Florida, and west along

33

Figure 27—Distribution of the Seminole bat in South Carolina (Menzel and others2003). 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 theSoutheastern United States


- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -Female


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

Mass 14 12.7 1.80 14.15 8.5 – 15.0

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -Male


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

Mass 12 9.8 1.18 12.09 7.8 – 11.0


RoostingIn the winter, Seminole bat roosts

commonly are found in oak hammockcommunities in Spanish moss(Constantine 1958), but the species’winter roosting habits in SouthCarolina are not known.

Seminole bats also commonly roostin Spanish moss during the summermonths (Barbour and Davis 1969).Although one summer roost waslocated under loose bark, this speciestypically does not use bark roosts(Sealander 1979). On the SavannahRiver Site, summer roosts usually arein the terminal branches of pine limbsin communities dominated by pines(Menzel and others 1998). Although 10tree species were used as roosts, most(86 percent) were pines, primarilyloblolly (Menzel and others 2000b).Roosts tended to be in taller, largerdiameter trees than the surroundingoverstory (Menzel and others 2000b).Roosts were found in areas with higherbasal area, lower understory speciesrichness, and less Spanish moss thanrandom plots (Menzel and others2000b). On-site, Seminole bats move


the Gulf Coast into east Texas (Wilkins1987). Museum collections include 42Seminole bats from South Carolina; 58have been live-captured in the State.Distribution of the Seminole bat inSouth Carolina includes the upper andlower Coastal Plain (fig. 27). They havebeen captured in the Piedmont andBlue Ridge, but their distribution insurrounding States suggests theyrarely occur above the fall line.

The Seminole bat was one of themost commonly captured bat speciesduring the 1979 survey (Anon. 1980).In surveys conducted between 1996and 2001, we captured six adultfemales, seven adult males, six juvenilefemales, and three juvenile males(table 7). Based on surveys conductedon the Site, the Seminole bat is thethird most common species, exceededin abundance only by evening andeastern red bats.


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Figure 28—Hoary bat.

to new roost trees often, spendingan average of 1.7 days in each. Onaverage, they select new roost treeswithin about a 0.25-ha portion of theirhome range.

Foraging and Home RangeAlthough Seminole bats commonly

forage at treetop level (Barbour andDavis 1969), two reports documentthe species gleaning prey fromleaf surfaces (Barbour and Davis1969, Sherman 1935). Sherman(1939) found the remains ofHomopterans (Jassidae), Dipterans(Dolichopodidae, Muscidae), andColeopterans (Scolytidae) in a singlestomach. Zinn (1977) reportedthat Coleoptera, Odonata, andHymenoptera were importantfood items. Carter (1998) foundthat Seminole bats fed primarilyon Hymenoptera, Coleoptera, andLepidoptera on the Site.

Radio tracking on the Site indicatedthe home-range size of five Seminolebats to be between 189.2 and 704.4 ha,

with an average of 423.8 ha (Carter1998). Habitat types in the home-rangeareas included pine forests (55percent), bottomland hardwoods(35 percent), and upland hardwoods(11 percent) (Carter 1998).


Because it is hard to differentiatecalls of eastern red and Seminole bats,we included both calls in a singlecategory. So far, using acousticalmonitoring to determine the effect ofhabitat type and stand age on the flightactivity of this species has not beenpossible. See the eastern red batspecies account for a description of theeffect of habitat type and stand age onthe flight activity of the eastern red/Seminole bat group.

ReproductionPregnant Seminole bats have been

collected between early May and mid-June (Barkalow 1948, Coleman 1950,Jennings 1958, Moore 1949).

Parturition usually occurs in June andJuly, but the species’ reproductivehabits in South Carolina are notknown. However, we captured juvenileSeminole bats on the Savannah RiverSite and tracked adult females toroosts occupied by their young.Seminole bats raise their progenyon the Site.

Hoary bat(Lasiurus cinereus)

Morphology and DistributionThe hoary bat is the largest bat

species found in South Carolina. It hasfrosted pelage and cream-colored earswith 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 massranges from 18 to 38 g, and its averagetotal length is 134.8 mm (Whitakerand Hamilton 1998). See table 15 foraverage body measurements ofindividuals collected in the Southeast.

35

Figure 29—Distribution of the hoary bat in South Carolina (Menzel and others2003). Symbols denote counties for which records exist.

Hoary bat(Lasiurus cinereus)

Table 15—Body measurements of the hoary bat in the SoutheasternUnited States


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


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

Mass 6 21.2 2.40 11.32 19.0 – 24.9


The hoary bat has a more extensivedistribution than any other bat speciesin South Carolina. Its range extendssouth from northern Canada near thetree line through Central America. Thesouthern limit of its range extends intoArgentina and Chile (Shump andShump 1982b). Museum collectionsinclude two hoary bats from SouthCarolina; three have been live-capturedin the State. The species’ distributionin South Carolina (fig. 29) includes allfour physiographic provinces, and itsstatus is listed as undetermined (table9) (South Carolina Department ofNatural Resources 2001). Because ofits migratory patterns, it probably israre in the State during summer.

We captured three hoary bats (twofemales and one sex undetermined) onthe Site from 1996 to 2001 (table 7).Whitaker and Hamilton’s (1998)summer range map for hoary batsdoes not include South Carolina.Although our surveys indicate thatthey do occur on the Site in summer,they are not abundant.

RoostingAlthough the hoary bat’s winter

roosts typically are found in treefoliage, they also have been found intree cavities and squirrel nests(Constantine 1966, Cowan and Guiguet1965, Neill 1952). The species typicallydoes not roost in caves (Mumford1953, Myers 1960).

Little is known about the hoary bat’ssummer or winter roosting habits inSouth Carolina, and nothing is knownabout its roosting requirements onthe Site.

Foraging and Home RangeHoary bats begin foraging in late

evening (Barbour and Davis 1969),sometimes establishing foragingterritories (Barclay 1984a). Prey itemsinclude Coleopterans, Dipterans,Orthopterans, Isopterans, Odonatans,Hymenopterans, and Lepidopterans(Black 1972, 1974; Ross 1967;Whitaker 1972; Whitaker and others1977), but it seems to preferLepidopterans (Black 1972, Ross1967). Its foraging habits in South



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

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Figure 31—Big brown bat.Ph

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Carolina are poorly understood.However, its diet on the SavannahRiver Site probably is similar to its dietelsewhere in its range.

We have found no information aboutthe size or spatial characteristics of thespecies’ home range.


We recorded hoary bats on 18.8percent of the lake and pond surveypoints, 5.7 percent of the Carolina baysurvey points, 7.6 percent of thebottomland hardwood survey points,2.6 percent of the forest gap surveylocations, 6.8 percent of the grass-brush survey points, none of theupland hardwood points, none of thepine-hardwood points, 5.0 percent ofthe loblolly-slash survey points, and1.4 percent of the longleaf surveylocations. Overall, we recorded it on5.2 percent of all survey locations(fig. 30). Its activity level waslow throughout the Site, primarilyconcentrated in bottomland hardwoodstands (table 4).

Stand age affected hoary bat activitydifferently than it affected that of otherbat species on the Site (fig. 9). Basedon the low call frequency, highwing loading, and moderately highwing-aspect ratio of the species, weexpected activity would be highest inclearcuts; yet we recorded no hoarybat calls there and found most activityconcentrated in stands aged > 60years (table 5).

ReproductionHoary bats probably mate during

autumn migration (Shump and Shump1982b). Parturition occurs from Maythrough July (Kurta 1982, McClure1942, Mumford 1969, Provost andKirkpatrick 1952, Whitaker andMumford 1972). Because the speciesmigrates north in spring, parturitionmay not occur in South Carolina.

Big brown bat(Eptesicus fuscus)

Morphology and DistributionThe big brown bat is a medium

sized, nondescript brown bat and isthe third largest in South Carolina (fig.31). One of the 11 subspecies of bigbrown bat, E. f. fuscus, occurs in theState (Kurta and Baker 1990). Its massranges from 13 to 25 g, and its averagetotal length is 114.3 mm (Whitaker and

Hamilton 1998). See table 16 foraverage body measurements ofindividuals collected in the Southeast.

The range of the big brown batextends from the Atlantic to the PacificCoasts, and south from Canadathrough Central America and intoBrazil (Kurta and Baker 1990). It isfound throughout the continentalUnited States and is the most commonbat species throughout much of itsrange. It is most abundant in the



Figure 32—Distribution of the big brown bat in South Carolina (Menzel andothers 2003). Symbols denote counties for which records exist.

Big brown bat(Eptesicus fuscus)

Table 16—Body measurements of the big brown bat in theSoutheastern United States


- - - - - - - - - - - - - - mm - - - - - - - - - - - - - -Female


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

Mass 21 17.2 4.04 23.54 11.0 – 27.0

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

MaleTotal 16 107.6 6.59 6.13 97.0 – 119.0Tail 16 41.7 4.67 11.18 30.0 – 49.0Foot 16 9.7 1.08 11.13 8.0 – 12.0Ear 15 16.4 1.80 11.00 13.0 – 19.0Tragus 14 6.4 1.11 17.40 5.0 – 8.0Forearm 15 46.6 2.53 5.43 40.0 – 50.0

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

Mass 16 12.8 3.12 24.33 7.0 – 18.8


39

deciduous forest biome (Kurta andothers 1989). Museum collectionsinclude 15 big brown bats from SouthCarolina; 18 have been live-capturedthere. It is common throughout theState (fig. 32) and is found in all fourphysiographic provinces.

Despite its apparent abundance inSouth Carolina, this species was notdocumented in the 1979 survey, andbetween 1996 and 2001 we capturedonly six (two adult females, threeadult males, and one juvenile female)(table 7). The relatively few captureson the Site probably was due to thescarcity of preferred roosts. Althoughit uses many different structures(see Roosting section), summerroosts are most often found in theattics of houses.

RoostingBig brown bats are one of the most

studied in North America. Theirhibernacula have been found inmines, caves, rock crevices, stormsewers, and buildings (Barbourand Davis 1969, Beer 1955, Fenton1972, Mills and others 1975, Mumford1958, Nagorsen 1980, Phillips 1966,Quay 1949).

Summer roosts usually are found inhollow oak and American beech(Fagus grandifolia Ehrh.), but mostcommonly the species uses human-made structures such as barns orhouses (Barbour and Davis 1969,Christian 1956, Kurta 1982). In earlyAugust 1996, Menzel (1998) andCarter (1998) radio tracked two bigbrown bats in a bottomland hardwoodswamp to a maternity roost in a hollowbald cypress snag along the SavannahRiver. Other than this anecdotalaccount, no studies have investigated

the species’ summer or winter roostinghabits in South Carolina.

Foraging and Home RangeBig brown bats typically begin

foraging within the first hour aftersunset (Kurta 1982, Phillips 1966).They feed on Isoptera, Hemiptera,Homoptera, Hymenoptera,Lepidoptera, and Diptera (Freeman1981, Menzel and others 2000a, Ross1967), although most of their diet iscomposed of beetles (Hamilton 1933;Menzel and others 2000a; Phillips1966; Whitaker 1972, 1995). Menzeland others (2000a) reported thatduring the reproductive period,females selectively forage onColeopterans and eat fewerLepidopterans, Dipterans, andHymenopterans, based on theavailability of such insect ordersin the foraging area. The species’foraging habits in South Carolinaare not known.

To date, little quantitativeinformation has been gathered aboutthis species’ foraging habitat selection.Furlonger and others (1987) found thatthey were generalists when selectinghabitat. There are many reports offoraging around urban streetlights,e.g., Geggie and Fenton 1985. In theGeorgia Piedmont, Menzel and others(2001c) found that big brown batsselectively foraged in rural rather thanurban areas, and less often inagricultural fields and clearcuts than inhardwood and pine forests. Menzeland others (2001c) reported that theaverage home-range size was 2906 ha;Brigham (1991) reported theirmaximum foraging distance to be 4.4km. Both these sets of data suggestlarge foraging ranges. Informationabout the foraging behavior of this

species in other regions of its rangesuggests that foraging activity on theSite would not be concentrated in asingle habitat type, but would bedispersed across a variety of types.The species probably forages inboth upland and bottomland habitatson the Savannah River Site.


Big brown bats were recorded at25.0 percent of the lake and pondsurvey points, 20.0 percent of theCarolina bay survey points, 15.2percent of the bottomland hardwoodsurvey points, 5.3 percent of the forestgap survey locations, 18.6 percent ofthe grass-brush survey points, noneof the upland hardwood points, 7.1percent of the pine-hardwood points,7.9 percent of the loblolly-slash surveypoints, and 10.0 percent of the longleafsurvey locations. We recorded them on11.8 percent of all survey locations (fig.33). Activity was concentrated overlakes and ponds, grass-brush, and inbottomland hardwoods (table 4).Foraging activity appeared to beunaffected by stand age (fig. 9).

ReproductionMating occurs in September and

March (Mumford 1958, Phillips1966), and fertilization in spring(Wimsatt 1944). The species’reproductive habits are unknown inSouth Carolina, but we captured ajuvenile on the Site and tracked twofemales to a large (> 30 individuals)maternity colony located in a baldcypress snag where Pen Branch flowsinto a swamp forest adjacent to theSavannah River. The species probablyraises its young on-site.



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.

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Figure 34—Little brown bat.

Table 17—Body measurements of the little brown bat in theSoutheastern United States


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


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

Mass 10 6.5 1.09 16.62 5.5 – 9.3


Bats of SouthCarolina in AreasNeighboring the

Savannah River Site

FAMILY VESPERTILIONIDAE

Little brown bat(Myotis lucifugus)

Morphology and Distribution

T he little brown bat is a smallbrown bat with glossy pelagethat occurs in some areas of

South Carolina (fig. 34). Of the sixsubspecies currently recognized, onlyM. l. lucifugus occurs in the State(Fenton and Barclay 1980). Its massranges from 4 to 8 g, and its averagetotal length is 89.2 mm (Whitaker andHamilton 1998). See table 17 foraverage body measurements ofindividuals collected in the Southeast.

The little brown bat is commonthroughout much of the United States.In the Eastern States, the southernlimit of its range reaches into northernportions of South Carolina, Georgia,Alabama, and Mississippi (Fentonand Barclay 1980). Museumcollections include three in South

Carolina; three have been live-capturedthere. Although its typical distributionin the State is limited to the BlueRidge (fig. 35), captures have beenconfirmed in Beaufort County onthe lower Coastal Plain (Davis andRippy 1968). South Carolina has listedit as uncommon or rare (South

Bats of South Carolina in Areas Neighboring the Savannah River Site


Figure 35—Distribution of the little brown bat in South Carolina (Menzel andothers 2003). Symbols denote counties for which records exist. Records south of

the range line likely are extralimital, because the primary distribution of thisspecies is in the Blue Ridge and northward.

Table 18—Body measurements of the eastern small-footed myotis inthe Southeastern United States


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

Total 2 78.0 2.83 3.63 76.0 – 80.0Tail 2 33.0 1.41 4.28 32.0 – 34.0Foot 2 6.0 1.41 23.57 5.0 – 7.0Ear 2 14.0 0. 0. 14.0 – 14.0Tragus 2 5.0 NA NA 5.0 – 5.0Forearm 1 31.0 1.41 4.56 30.0 – 32.0

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

Mass 2 4.2 0.07 1.66 4.2 – 4.3


Carolina Department of NaturalResources 2001).

One female little brown bat wascaptured on the Site in 1996 (table 7).Because the Site is south of the species’typical range, that single recordprobably is extralimital. The speciesprobably does not occurthere regularly.

RoostingLittle is known about the roosting

habits of little brown bats in SouthCarolina. Its hibernacula mostcommonly are found in caves andmines (Baker 1965, Humphrey andCope 1976), but no hibernacula havebeen found in the State. In summer,little brown bats roost under rocks, inpiles of wood, and in trees (Fenton andBarclay 1980). Maternity roostscommonly are found in buildings(Davis and Hitchcock 1965, Griffin1940, Youngman 1975). The summeror winter roosting habits of thisspecies in South Carolina have notbeen reported.

Foraging and Home RangeLittle brown bats emerge shortly

after dusk to feed (Nagorsen andBrigham 1990). Numerous foraginghabit studies have been conducted(Anthony and Kunz 1977; Belwood andFenton 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 andothers (1960) found that when feedingin an insect swarm, little brown batscould capture up to 12 fruit flies perminute. Although the species’ diet isvariable, aquatic insects constitutemost of its forage (Fenton and Barclay1980). Little is known about the size ofits home range, and nothing is knownabout its home range or habitat use inSouth Carolina. Because the speciesdoes not occur there regularly, itsforaging behavior should be of littleconcern to land managers on the Site.


We did not record any calls oflittle brown bats during the 2001AnaBat survey.

ReproductionLittle brown bats mate in autumn,

and fertilization occurs in spring whenfemales leave the hibernacula.Duration of hibernation typicallydecreases with decreasing latitude,resulting in earlier ovulation andparturition dates at more southerlylatitudes (Fenton and Barclay 1980).Parturition occurs in late May andJune (Barbour and Davis 1969,Hayward 1963, Hoffmeister 1989)on the Savannah River Site, and it isunlikely that little brown bats raisetheir young there.

43

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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 DistributionThe eastern small-footed myotis

is a small brown bat with long, glossypelage and a black mask (fig. 36).It is the smallest bat found in SouthCarolina. This species is monotypic(van Zyll de Jong 1985). Its massranges from 4 to 6 g, and its averagetotal length is 77.5 mm (Whitakerand Hamilton 1998). See table 18for average body measurements ofindividuals collected in the Southeast.

The species’ range is restricted tonortheastern North America, fromextreme northwestern South Carolinawest into Oklahoma and Missouri andnorth into the St. Lawrence forestregion of Ontario and Quebec (van Zyllde Jong 1985). Museum collectionsinclude 42 eastern small-footed myotisfrom South Carolina. Within the State,its range is limited to the Blue Ridgein the extreme north (fig. 37). Thespecies appears to be uncommon



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Figure 38—Northern long-eared myotis.

throughout its range, and it is listed asthreatened in South Carolina (table 9)(South Carolina Department of NaturalResources 2001).

No eastern small-footed myotis’ havebeen captured on the Site. Based onits distribution and roosting-habitatpreferences, i.e., rock outcrops, it ishighly unlikely that this species wouldoccur on the Site.

RoostingRoosting habits of eastern small-

footed myotis’ in South Carolina arenot well known. In other States,hibernacula have been found in cavesand mines (Davis 1955; Fenton 1972;Hitchcock 1945, 1949; Mohr 1936).Summer roosts have been foundunder rocks and in buildings (Barbourand Davis 1969, Hitchcock 1955,Tuttle 1964).

Foraging and Home RangeEastern small-footed myotis’ begin

foraging shortly after sunset. Average

foraging height is 1 to 3 m above theground (van Zyll de Jong 1985), butlittle is known about its prey.

Because it is so small, this species isdifficult to track using radiotelemetry.To lessen the transmitter’s effect onforaging behavior, transmitter masswas < 0.4 g. Because of this difficulty,and because of the species’ rarity,characteristics of its home range havenot been documented.


We did not record any eastern small-footed myotis calls during the 2001AnaBat survey. This species has notbeen captured on the Site, and it isunlikely to occur there.

ReproductionInformation about eastern small-

footed myotis reproduction is limited toone record of a maternity colony locatedbehind the door of a barn in Ontario,Canada, in summer (Hitchcock 1955).

No maternity colonies have beenfound in South Carolina.

Northern long-eared myotis(Myotis septentrionalis)

Morphology and DistributionThe northern long-eared myotis is

a medium-sized bat with long, glossybrown pelage (fig. 38) (Fitch andShump 1979). Its mass ranges from5 to 10 g, and its average total lengthis 84.1 mm (Whitaker and Hamilton1998). See table 19 for average bodymeasurements of individuals collectedin the Southeast.

The northern long-eared myotisoccurs throughout much of the easternhalf of the United States. Its rangeextends east from Saskatchewanto Quebec, Canada, and south toFlorida (Fitch and Shump 1979).Although there are no museumspecimens from South Carolina, 22individuals have been live-captured insurveys. 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 inthe Southeastern United States


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


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

Mass 12 8.0 11.65 144.92 3.6 – 45.0


restricted to the Blue Ridge, where thespecies is common (fig. 39) (SouthCarolina Department of NaturalResources 2001).

No northern long-eared myotishave been captured on the SavannahRiver Site. Its typical range does not

include the Site, and it is unlikelyany occur there.

RoostingHibernacula used by northern long-

eared myotis have been found in caves(Griffin 1940, Swanson and Evans1936), mines (Stones and Fritz 1969),

and a storm sewer (Goehring 1954).During summer, the species roostsin buildings (Doutt and others 1966,Turner 1974), behind shutters(Mumford 1969), and under the barkand in the cavities of trees (Menzeland others 2002d, Mumford andCope 1964). Its summer or winterroosting habits in South Carolina arenot known.

Foraging and Home RangeNorthern long-eared myotis forage

shortly after dusk and before dawn(Barbour and Davis 1969). They foragearound ponds and along hillsides, bothabove (Cowan and Guiguet 1965) andbeneath (LaVal and others 1977) thetree canopy. Little is known about theirdiet. Their foraging habits in SouthCarolina are not known.

Although this species’ home-rangecharacteristics in the Southeasthave not been studied, data areavailable for the central Appalachians.In the Allegheny Mountains of WestVirginia, its home ranges areunimodal, from 43 to 578 ha (mean216 ha), and include clearcuts,deferment harvests, streams, roadcorridors, and mature second-growtheastern deciduous forests.

4

Effect of Stand Age andHabitat Type on Flight Activity

We did not record any calls ofnorthern long-eared myotis duringthe 2001 AnaBat survey. We have notcaptured any of this species on theSite, and it is unlikely to occur there.

ReproductionNorthern long-eared myotis copulate

in autumn, and sperm is stored inthe uterus throughout hibernation.Fertilization occurs in spring (Barbourand Davis 1969). Parturition occurs inJune, and the young typically becomevolant in late July (Easterla 1968, Kunz1971). Parturition times and maternitycolony habits in South Carolina arenot known.



Figure 40—Northern yellow bat.

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5 Menzel, M.A.; Krishon, D.M.; Carter, T.C. 1995.Roosting, foraging, and habitat use by bats ofSapelo Island, Georgia. Unpublished Tech. Rep.60 p. On file with: Georgia Department of NaturalResources, Attn: J. Ozier., Social Circle, GA.

Northern yellow bat(Lasiurus intermedius)

Morphology and DistributionThe northern yellow bat is a large

yellow bat and is the second largest batin South Carolina (fig. 40). Althoughtwo subspecies of it are recognized,the validity of this designation is amatter of debate (Whitaker andHamilton 1998). Of the two subspecies,L. intermedius floridanus occurs inSouth Carolina. Its mass ranges from14 to 20 g, and average total length is

126.8 mm (Whitaker and Hamilton1998). See table 20 for average bodymeasurements of individuals collectedin the Southeast.

Although one northern yellow batwas collected in New Jersey (Koopman1965), typically the species is restrictedto coastal areas of the SoutheasternUnited States and Central America(Webster and others 1980). Museumcollections include 11 northern yellowbats from South Carolina; one hasbeen live-captured in the State. Itoccurs in the lower Coastal Plain, and

its range extends into the upperCoastal Plain along the Savannah River(fig. 41). The State has listed the statusof this species as undetermined (table9) (South Carolina Department ofNatural Resources 2001).

One northern yellow bat wascollected along Upper Three RunsCreek in 1978. However, none wascaptured during the 1979 survey(Anon. 1980) or the 1996 to 2001survey. Based on the species’distribution, it probably is nota common resident on theSavannah River Site. The individualcollected along Upper Three RunsCreek probably represents anextralimital record.

RoostingLittle is known about the northern

yellow bat’s winter roosting habits inSouth Carolina or elsewhere.

Summer roosts have been locatedin palm groves (Davis 1960), onhardwood stems (Rageot 1955), inpine-oak woodlands (Carter and Jones1978, Carter and others 1966, Jones1964, Sherman 1944), and in Spanishmoss (Jennings 1958).

5 Northern

yellow bats usually roost alone;however, Baker and Dickerman (1956)reported seeing about 45 individualsfly from a communal roost under cornstalks hanging from the side of an oldtobacco curing shed. The speciesis not a regular summer resident onthe Site and should be of littlemanagement concern.

Foraging and Home RangeNorthern yellow bats typically begin

foraging well before dark (Lowery1974). They forage 4 to 6 m above theground (Barbour and Davis 1969) andprefer open foraging areas such asairports, golf courses, and fields(Jennings 1958). Lowery (1974)observed them foraging over theMississippi River. After the youngbecome volant, they form feeding

47

Figure 41—Distribution of the northern yellow bat in South Carolina (Menzel andothers 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 theSoutheastern United States


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


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

Mass 3 20.0 2.29 11.46 17.5 – 22.0


aggregations that may consist of> 100 individuals (Jennings 1958). Suchaggregations primarily are composedof females. Northern yellow batsemerge and feed on warm nightsthroughout winter (Jennings 1958).

Northern yellow bats feed onHomoptera, Zygotera, Diptera,Coleoptera, and Hymenoptera(Sherman 1939), but their foraginghabits in South Carolina arenot known.

The home range of a northernyellow bat tracked on Sapelo Island,GA, was 10.5 ha (Krishon and others1997). It was located in oak (73percent) and slash-loblolly pine (25percent) communities. The averagedistance from the roost area center to aforaging location was 109 m. This is arelatively small home range comparedto 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 oneindividual and may not represent thespecies’ average home-range size. Nohome range or habitat use studies ofnorthern yellow bats have beenconducted in South Carolina.


We did not record any calls ofnorthern yellow bats during the 2001AnaBat survey. Although we collectedone on the Site, it is unlikely that thisspecies regularly occurs there.

ReproductionParturition occurs in late May and

June (Lowery 1974, Schmidly 1991).Litter size ranges from two to fourindividuals and averages 3.4 per litter(Jennings 1958). Mating probablyoccurs in autumn and fertilization inspring (Lowery 1974). The species’reproductive habits in South Carolinaare not known.



Figure 42—Brazilian free-tailed bat.Ph

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FAMILY MOLOSSIDAE

There are 16 genera and 86 speciesin the family Molossidae (Nowak1994). Members of this family arefound in southern Europe, Asia, Africa,and from the northern half of SouthAmerica north into the United States.Some are colonial and form thelargest aggregations of mammalsdocumented. Davis and others (1962)estimated that summer molossidpopulations in four caves (Bracken,Goodrich, Rucker, and Frio) in Texasexceeded 10 million individuals duringpeak periods. Molossids are mediumto large bats. Because one of the mostdefining characteristics of this familyis that the tail extends beyond theposterior border of the uropatagium,members of this family commonly arecalled the free-tailed bats.

Two genera and six species ofmolossids occur in the United States.One species occurs in South Carolina.

Brazilian free-tailed bat(Tadarida brasiliensis)

Morphology and DistributionThe Brazilian free-tailed bat is the

only species of free-tailed bat thatoccurs in South Carolina (fig. 42).As its name suggests, the tail of thisspecies extends beyond the posterioredge of the uropatagium. It is thesmallest member of its genus in theUnited States. Of the nine subspeciesnow recognized, only T. brasiliensiscynocephala occurs in the State(Wilkins 1989). Its mass ranges from8 to 14 g, and its average total lengthis 91.9 mm (Whitaker and Hamilton1998). See table 21 for average bodymeasurements of individuals collectedin the Southeast.

The Brazilian free-tailed bat has oneof the largest ranges of any mammalin the Western Hemisphere (Wilkins1989). From South Carolina, thespecies’ range extends west intosouthern Oregon and south throughCentral America. It reaches the

southern limit of its range in thePatagonian region of southern Chileand Argentina (Wilkins 1989). A totalof 338 Brazilian free-tailed bats fromSouth Carolina are in museumcollections. Its distribution in the Stateextends through the upper and lowerCoastal Plain (fig. 43). There havebeen live captures in the Piedmont,but these isolated records may notreflect the species distribution inSouth Carolina.

Free-tailed bats have been submittedfor rabies testing from Aiken andBarnwell Counties, SC. Although nonewas captured during the 1979 or 1996to 2001 surveys, it was recordedduring a 1996 AnaBat study onSavannah River Site (Menzel andothers 2002d). Additionally, its rangein the Southeast, the individualssubmitted for rabies testing fromAiken and Barnwell Counties, andrecent reports that it is roosting in

49

Figure 43—Distribution of the Brazilian free-tailed bat in South Carolina (Menzeland others 2003). Symbols denote counties for which records exist.

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


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


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

Mass 12 11.5 2.03 17.27 9.3 – 16.0


structures near Clemson, SC, suggestthat it may be more common on the Sitethan survey results indicate.

6 Brazilian

free-tailed bats typically roost in human-made structures. Their dependence onsuch structures for roosting habitat maylimit their abundance on the Site.

RoostingThere is little difference between the

summer and winter roosting habits ofBrazilian free-tailed bats, although theydo vary among different regions of theUnited States. Most roosts in theSouthwest are found in caves, althoughthe bat does use buildings sometimesduring migration. In the Southeast,roosts typically are found in buildings(Barbour and Davis 1969), but theyalso have been found in hollow treesand in the expansion joints of bridges(Lowery 1974, Tuttle 1994). Thisspecies is very gregarious and almostalways roosts in colonies (Barbour andDavis 1969). Although the size of cavecolonies of the Southwest commonlyexceeds 1 million individuals, coloniesin the Southeast typically arecomposed of < 50,000. Roosting habitsin South Carolina are not known.Although nothing is known about thespecies’ on-site roosting requirements,there is no reason to suspect thatthey are different from other areasof the Southeast.

Foraging and Home RangeAlthough emergence time is

variable, Brazilian free-tailed batsusually leave their roost shortly aftersunset (Bailey 1951). As with most batspecies inhabiting temperate regions,the length of the foraging bout varieswith temperature. Bailey (1951) foundthat on warm nights, individuals in acolony of Brazilian free-tailed bats inLouisiana did not return to the roostuntil morning. Most studies of thisbat’s foraging habits have beenconducted in the Southwest. In theSoutheast, Brazilian free-tailed batsfeed on Hymenoptera, Coleoptera,Diptera, and Lepidoptera (Sherman

6 Personal communication. 1999. Mary Bunch,Wildlife Biologist, South Carolina Department ofNatural Resources, Wildlife Diversity Section,P.O. Box 1806, Clemson, SC 29633.



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

oto

cour

tesy

of J

ohn

Mac

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gor

1939). Their diet in South Carolinaor on the Savannah River Site has notbeen studied. There is no reason tosuspect that it would be any differentthan elsewhere in the Southeast, butbecause little is known about thisspecies in the region, and becauseits diet may differ from the westernsubspecies, its diet in the easternUnited States warrants future study.

Nothing is known about the sizeof the Brazilian free-tailed bat’s homerange in the Southeast; further studywill be necessary to determine thespatial extent of its foraging areas inthe region.


We did not record any calls ofBrazilian free-tailed bats during theAnaBat survey. However, the specieswas detected during call surveysconducted in canopy gaps in abottomland hardwood forest duringsummer 1997 (Menzel and others2002a). Brazilian free-tailed batshave been collected from Aiken andBarnwell Counties, SC, and fromareas south and north of the Site.

ReproductionIn Florida, spermatogenesis occurs

throughout late autumn and winter(Sherman 1937). Unlike vespertilionidbats in South Carolina, which mate inautumn and ovulate in spring, Brazilianfree-tailed bats both copulate andovulate in spring (Cockrum 1955).Breeding occurs in February andMarch (Golley 1966). Parturitionoccurs from late May to early August(Barbour and Davis 1969, Sherman1937), and the typical litter size is one(Wilkins 1989). Maternity coloniesof this species have been located inthe State, and the species occasionallymay raise its young on the SavannahRiver Site.

Conservation andResearch Needs

Although no federally listed batspecies occur on the Savannah River Site, the State of South

Carolina lists the Rafinesque’s big-eared bat as endangered and thesoutheastern myotis as threatened(table 9). Two of the nine species(22 percent) that occur on the Siteregularly are protected. In the

Southeast, 87 percent of all batspecies carry special conservationdesignations somewhere within theirregional distribution (Laerm andothers 2000). Worldwide, a relativelyhigh proportion of bat species aredesignated as threatened orendangered. All 15 bat speciesthat occur in Great Britain arenow considered threatened orendangered (Nagorsen and Brigham1990, Stebbings 1988), and bat

51

populations seem to be decliningglobally (Nowak 1994).

Potential issues of conservationconcern have been raised for the Siteand surrounding areas. More than 75percent of the Site is included in thetimber management program and istherefore subject to some timberharvesting (Blake, in press). Althoughthere has been a recent surge ofinformation concerning the effects offorest management practices on forestbat communities, e.g., Barclay andBrigham 1996, Krusic and others1996, little is known about the effectsof most silvicultural practices in theSoutheast. Studies investigating theeffects that harvesting techniques,e.g., clearcutting, group-selection cuts,and various intermediate treatments,e.g., thinnings, herbicide application,and prescribed burning, may have onbat communities will be necessaryto make more informed landmanagement decisions. As moreland area is devoted to short-rotationforestry practices, roosting habitatmay become scarce for bat speciesthat depend on older trees for cavity orbark roosts. Studies should investigatethe potential use of bat houses toenhance roosting habitat where ithas been compromised (fig. 44).

In addition to timber management,other large-scale agricultural practicesare common throughout much ofSouth Carolina. All nine speciesof bats that occur on the Site areinsectivorous and therefore may beaffected by pesticides that are appliedto agricultural crops such as cotton,soybeans, and corn. We know littleabout how these pesticides will affectbat populations over the long term.The effects of pesticides oncavernicolous species such as thegray bat have been examined(Clawson and Clark 1989), but theeffects of pesticides on tree-roostingbats are not known. All bat species thatoccur on the Savannah River Site roostin trees during at least part of the year.

Another conservation concern in theCoastal Plain of South Carolina focuseson the use of bridges as roosts by bats.In some areas of the United States,

such as Texas, bridges are importantroosting sites for many bat species(Keeley 1997). The endangeredRafinesque’s big-eared bat and thethreatened southeastern myotis roostunder bridges (Lance and others2001). Although many bridges on theSite may serve as roosts for thesespecies, no surveys have beenconducted to document the bats’use of them.

In the 1990s, great strides weremade towards understanding theroosting and foraging ecology of bats,but much remains unknown. Fewstudies have been conducted in theSoutheast, and results from studiesconducted in other regions of theUnited States and Canada may notbe applicable in South Carolina oron the Savannah River Site. To moreeffectively manage the Site’s batcommunities we need moreinformation about their roostingand foraging ecology.

Bats and Rabies

No work on bats is completewithout a discussion of rabies,a zoonotic disease that

generates considerable concernamong public health officials and analmost irrational fear among thegeneral public. Found worldwideexcept in parts of Scandinavia, theAustralian continent, and on islandmasses such as Hawaii, New Zealand,Papua New Guinea, Japan, and Taiwan,rabies is caused by exposure to andinfection by single-strand RNA virusesin the genus Lyssavirus, familyRhabdoviridae (Krebs and others1995, Wunner and others 1988). TheBritish Isles had long been thoughtto be rabies-free until the discoveryof a rabid Daubenton’s bat (Myotisdaubentonii) in 1996 (Whitby andothers 1996). Transmission to humansor other mammals occurs primarilyby introduction of the virus intowounds, cuts, scratches, and mucousmembranes from an infected animal,usually by biting (Macinnes 1987).Other routes of viral transmission,such as inhalation of rabies-infectedaerosol, are known, but they are

extremely rare (Constantine 1962,1967b; Winkler and others 1973), as istransmission through rabies-infectedhuman tissue transplanted inuninfected organ recipients (Centersfor Disease Control and Prevention1980, 1981).

Rabies was first documented in batsin the United States in 1953 (Krebs andothers 1994). Contrary to popularbelief, bats are not nonsymptomaticcarriers of rabies. However, the lengthof latency prior to onset of symptomsand eventual death in bats is highlyvariable (Bell and others 1969,Macinnes 1987, Moreno and Baer1980). The rabies virus is reservoiredin many wild mammal hosts, includingbats (Davidson and Nettles 1997),although transmission cycles usuallyare separate between terrestrialmammals and bats (Constantine 1967a,1979a; Smith 1989). In Latin America,vampire bats routinely spread rabiesto domestic livestock and humans(Arellano-Sota 1988, Batista-da-Costaand others 1993, Martinez-Burnes andothers 1997), and in North America,cases of rabies in terrestrial wildmammals and humans have resultedfrom contact with several of theinsectivorous bat species commonlyfound in South Carolina (Constantine1979b). Previous research linked theincidence of rabies in foxes to anumber of bat caves in the Karstlimestone regions of Tennessee(Fredrickson and Thomas 1965),but such relationships were poorlysubstantiated (Fischman 1976).Following the discovery of rabies virusstrains and species-specific variants,most supposed links between batrabies and the maintenance of rabiesin terrestrial mammals have beendiscounted (Krebs and others 1995,Smith 1989, Smith and others 1986).However, in areas where the incidenceof rabies in terrestrial mammals is low,bats are responsible for small rabiesoutbreaks or “clusters” (Daoust andothers 1996).

Among terrestrial mammals in SouthCarolina and throughout the Southeastand mid-Atlantic regions, raccoons(Procyon lotor) are important

Conservation and Research Needs ° Bats and Rabies


terrestrial reservoir species (Rosatteand others 1997). Skunks (Memphitismemphitis, Spilogale putorius) arerabies vectors in the Midwest andMississippi Valley (Krebs and others1994, 1995; Smith and others 1986),and foxes (Urocyon cinereoargenteus,Vulpes vulpes) are rabies reservoirsin Canada, New England, and parts ofthe Southwest (Davidson and Nettles1997, Smith and others 1986). Coyotes(Canis latrans) also appear to beimportant vectors throughout muchof the Southwest (Clark and others1994, Davidson and Nettles 1997).

In developing countries, humancases of rabies are relatively common,accounting for approximately 50,000deaths annually (Fishbein and Bernard1995). Most routes of human exposureto rabies are from unvaccinated dogsand cats, particularly in developingcountries, but wild animals, includingbats, are responsible for many humancases (Krebs and others 1995).Advances in public health—such asanimal control laws, mandatoryvaccination measures for dogs in manyjurisdictions, and aggressive andprompt medical attention followinganimal bites—have served to limithuman exposure to and infection withrabies in North America (Rosatte andothers 1997, Tierkel 1975). Today,human exposure to rabies in theUnited States is almost wholly limitedto contact from wild animals, of whichexposure to infected bats accounts forthe majority of cases documentedsince the 1980s (Childs and others1994, Hoff and others 1993, Krebs andothers 1995). Nonetheless, the overallincidence of rabies contracted frombats in North America is low (Krebsand others 1995, Rotz and others 1998,Tuttle and Kern 1981). Efforts todevelop effective oral rabies vaccinesfor wild, terrestrial mammals—and todistribute such vaccines by widespreadbaiting—are underway in the UnitedStates and Canada (Davidson andNettles 1997), but such preventivemeasures for bats have not beencontemplated.

In humans, rabies is almost alwaysfatal. Following exposure, the

incubation period may range from< 10 days to > 6 years (extremelatency). Commonly, incubation timefrom exposure to the first symptoms is30 to 90 days. The rabies virus infectsnervous tissue, eventually spreading tothe spinal cord and brain, producingencephalitic lesions. It moves thenthroughout the central nervous systemand to peripheral sites such as thesalivary glands, where it is shed insaliva (thus facilitating transfer toanother animal). Initial symptoms inhumans can include pain and burningnear the site of the wound whereexposure occurred, headache, fever,flu-like malaise, and mental changessuch as anxiety and apprehension. Asthe disease progresses, mentaldeterioration occurs in the form ofdisorientation and hallucinations.Aggressive behavior and hydrophobia,or “fear of water,” does occur in humancases, although in less than one-half ofdocumented rabies cases. Physically,difficulty in swallowing, musclespasms, and paralysis occur as thedisease advances; death usually resultsfrom respiratory failure. From theonset of initial symptoms, death mayoccur within 1 to a few weeks(Fishbein 1991, Jackson and others2003, Krebs and others 1995).

In wild mammals, rabies takes twoforms—a furious or aggressive formand a withdrawn or dumb form.Skunks, foxes, dogs, and raccoonsoften lose all fear of humans and otheranimals when exhibiting the furiousform of the disease, and they mayexhibit extremely aggressivebehaviors. Activity patterns maychange. For example, nocturnalanimals may become active duringthe day (Davidson and Nettles 1997,Macinnes 1987). Rabid bats may beirritable and bite conspecifics, butfurious behaviors, such as are seen inwild carnivores, are rare (Macinnes1987, Sulkin and Allen 1974, Tuttle andKern 1981). The extent to which bat-to-bat rabies transmission occurs throughcontact from grooming and nursing orfrom antagonistic interspecificencounters is unknown (Krebs andothers 1995, Macinnes 1987). Actualunprovoked attacks by rabid,

insectivorous bats on humans orother animals are extremely rare(Constantine 1979a, 1979b; Tuttle andKern 1981). Animals infected withrabies from bats often exhibit acreeping paralysis (Baer 1975). In thedumb form of rabies, animals remaincalm, yet are apparently disorientedand confused (Davidson and Nettles1997, Macinnes 1987). In most wildmammals, symptoms similar to thoseaffecting humans occur as the diseaseprogresses, advancing to paralysis andeventual death.

Following a suspected or confirmedexposure to rabies—via the use of afluorescent antibody test on braintissue of the suspect animal (Kissling1975, Wachendorfer and others1985)—development of rabies can beprevented by prompt postexposureprophylaxis of human antirabiesimmunoglobulin injections. Suchtreatment is followed by a series ofhuman diploid cell injections, rabies-adsorbed intramuscular vaccinations,or purified chick embryo cell culturevaccinations (Blythe and others 1998).For veterinarians, animal controlofficers, wildlife biologists, wildliferehabilitators, and zoologists, whocome into contact with wild and feralmammals on a regular basis,preexposure prophylaxis isrecommended (Krebs and others1995). Similarly, anyone handling batsor coming in close contact with batsshould be preimmunized.

For many years, rabies was thoughtto be a single viral type. With theadvent of monoclonal antibody surveysand advances in genetics researchthrough polymerase chain reaction(PCR) techniques, epidemiologistshave documented a number ofantigenically and genetically distinctvariants or strains of rabies virus(Koprowski and others 1985; Smithand others 1984, 1986). Strains uniqueto several species of bats throughoutNorth America have been documented(Smith 1988), and within a speciesthere may be more than one strain orvariant across wide geographic regionsor within a localized area (Smith andothers 1984). Thus, rabies strains

53

found in bats are distinct from thosein terrestrial mammals. As a result,public health officials and scientistscan differentiate and document routesof human exposure by animal typeand species. For example, of the 14confirmed cases of human rabiesin the United States contracteddomestically from 1980 to 1994, 11were linked to bats. Of those, eightwere attributed to viral strainsassociated with silver-haired bats(Krebs and others 1995), although thatspecies was not always the confirmedvector (Morimoto and others 1996),nor was evidence of a bat bite alwayspresent. Common carriers of the silver-haired bat rabies viral strain appear tobe big brown bats, little brown bats,and eastern pipistrelles (Messengerand others 1997).

Rabies has been documented inmost bat species that occur in SouthCarolina, including hoary bat, red bat,yellow bat, Seminole bat, big brownbat, little brown bat, small-footed bat,southeastern bat, gray bat, Townsend’sbig-eared bat, eastern pipistrelle,evening bat, silver-haired bat, andBrazilian free-tailed bat. Rabies has notbeen documented in Rafinesque’s big-eared bat (Constantine 1979b). Thecommon species most likely to occurin and near human dwellings andactivity centers, such as the big brownbat, probably pose the greatest risk fortransmitting rabies to humans (Childsand others 1994). Eastern red bats,perhaps the most widespread andnumerous species in South Carolina,accounted for a large percentage ofrabid bats collected in Illinois (Burnett1989), while a high percentage ofrabies-positive bats in Florida havebeen yellow bats (Bigler and others1975). Nationwide, the Centers forDisease Control and Preventionstatistics indicate that approximately10 percent of all annually reported andconfirmed rabid animals are bats(Krebs and others 1995). For example,in 1996, there were 7,124 documentedcases of nonhuman rabies in theUnited States, of which 741 were bats(Krebs and others 1997). Still, theoverall human health risks posed byrabid bats are extremely small in South

Carolina. Simple preventive measuressuch as avoiding handling any bat andavoiding entry into caves, attics, andabandoned buildings that harborroosting bats—or preventing roostingbats from using human-occupieddwellings—would eliminate mostroutes of contact and potential rabiestransmission. Immediate postexposurerabies prophylaxis should proceed ifone is bitten by or comes into closecontact with a bat, such as wakingto find a bat present in occupiedsleeping quarters.

Acknowledgments

We thank the U.S. Departmentof Energy; the USDA ForestService-Savannah River,

Southern Research Station, andNortheastern Research Station; theNational Council of the Paper Industryfor Air and Stream Improvement; WestVirginia University Division of Forestry;and West Virginia University Arlen G.and Louise Stone Swiger Fellowship forsupporting this project. Funding fromthe U.S. Department of Energy wasprovided through the USDA ForestService-Savannah River underInteragency Agreement DE-IA09-76SR00056. This project would nothave been possible without theprofessional support of Drs. B.R.Chapman and J. Laerm. Drs. B.R.Chapman and S.B. Castleberry providedhelpful comments onthe manuscript. This work wasconceived and supported throughoutby Dr. J.I. Blake.

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Whitaker, J.O., Jr. 1995. Food of the big brownbat Eptesicus fuscus from maternity colonies inIndiana and Illinois. American MidlandNaturalist. 134: 346–360.

Whitaker, J.O., Jr.; Hamilton, W.J., Jr. 1998.Mammals of the Eastern United States.Ithaca, NY: Cornell University Press. 583 p.

Whitaker, J.O., Jr.; Maser, C.; Cross, S.P.1981. Food habits of eastern Oregon bats,based on stomach and scat analyses.Northwest Science. 55: 281–292.

Whitaker, J.O., Jr.; Maser, C.; Keller, L.E.1977. Food habits of bats of western Oregon.Northwest Science. 51: 46–55.

Whitaker, J.O., Jr.; Mumford, R.E. 1972.Notes on occurrence and reproduction of batsin Indiana. Proceedings of the IndianaAcademy of Science. 81: 376–383.

Whitaker, J.O., Jr.; Rissler, L.J. 1992.Seasonal activity of bats at Copperhead Cave.Proceedings of the Indiana Academy ofScience. 101: 127–134.

Literature Cited

Whitby, J.E.; Johnstone, P.; Parsons, G.[and others]. 1996. Ten–year survey ofBritish bats for the existence of rabies.Veterinary Review. 139: 491–493.

Wilkins, K.T. 1987. Lasiurus seminolus.Mammalian Species. 280: 1–5.

Wilkins, K.T. 1989. Tadarida brasiliensis.Mammalian Species. 331: 1–10.

Wimsatt, W.A. 1944. Notes on breedingbehavior, pregnancy, and parturition in somevespertilionid bats of the Eastern UnitedStates. Journal of Mammalogy. 25: 23–33.

Winkler, W.G.; Fashinell, T.R.; Leffingwell,L. [and others]. 1973. Airborne rabiestransmission in a laboratory worker. Journalof American Medical Association. 226:1219–1221.

Workman, S.W.; McLeod, K.W. 1990.Vegetation of the Savannah River Site: majorcommunity types. SRO–NERP–19. Aiken, SC:U.S. Department of Energy, Savannah RiverEcology Laboratory. 137 p.

Wunner, W.H.; Larson, J.K.; Dietzschold,B.; Smith, C.L. 1988. The molecular biologyof rabies viruses. Review of InfectiousDiseases. 10(Suppl. 4): S771–S784.

Yougman, P.M. 1975. Mammals of the YukonTerritory. National Museum of CanadaPublications in Zoology. 10: 1–92.

Zinn, T.L. 1977. Community ecology of Floridabats with emphasis on Myotis austroriparius.Gainesville, FL: University of Florida. 88 p.M.S. thesis.

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 34Southeastern myotis 2 1 3 3 3 1 3 3 38Little brown bat 2 1 3 3 3 1 3 3 38Evening bat 1 1 1 3 3 1 2 3 30Rafinesque’s big-eared bat 2 1 2 3 3 1 3 3 36Silver-haired bat 2 1 2 3 3 1 3 3 36Eastern red bat 1 1 2 3 3 1 2 3 32Seminole bat 1 1 2 3 3 1 2 3 32Northern yellow bat 1 1 1 3 3 1 2 3 30Hoary bat 1 1 2 3 3 1 2 3 32Big brown bat 2 1 1 3 3 1 2 3 32Brazilian 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 occasionallyoccurs there.

Appendix

Appendix

KEYS TO THE SKINS

AND SKULLS OF SOUTH

CAROLINA BATS

Bat research as well as batmonitoring in the Southeasthave been impeded by

difficulty in training field personnelto reliably identify captured bats.Although many published keys containinformation about particular species inthe region (Barbour and Davis 1969,Golley 1962, Hoffmeister 1989, Jenkins1949, Schmidly 1991, Whitaker andHamilton 1998), many of them havebeen unreliable tools for teachingstudents and field personnel to identifybats by their skins and skulls. Mostkeys require the lower jaw, are poorlyor inadequately illustrated, or containunreliable univariate discriminatorymeasurements. To improve fieldidentification capabilities we devised atool combining traits that have been

useful in the field and the lab withinformation from other publishedkeys. Using the combined keys hasfacilitated relatively easy and accurateidentification of the 14 bat speciesthat occur in South Carolina. Thekeys are reprinted from Menzel andothers (2002a).

We used measurements frommuseum specimens, informationcontained in recent publications aboutbat morphological characteristics,and information from publisheddichotomous keys (Barbour and Davis1969, Golley 1962, Hoffmeister 1989,Jenkins 1949, Schmidly 1991). Becausethe bat’s lower jaw often is damagedor missing from museum skeletalspecimens or skulls found in the field,we devised a key that does not requirethe lower jaw for identification.

The eastern red (Lasiurus borealis)and Seminole (L. seminolus) bats aretwo of the most common species onthe Site, but many keys to bats in theSoutheast do not differentiate betweentheir skulls. Because these two species

are relatively abundant on the Site, wewere able to incorporate informationabout the size of the protuberance ofthe lacrimal ridge, i.e., the lacrimalshelf, and accurately differentiateamong about 75 percent of easternred and Seminole bats skulls foundthere (Laerm and others 1999,Lowery 1974).

The key to the skulls allows a userto classify a specimen as belonging tothe genus Myotis. Because simpleunivariate measures cannot be usedto differentiate among southeasternmyotid skulls, identification to speciescannot be accomplished with this key.However, many nonmyotid bat skullscan be identified to species bycounting the number of teeth in oneupper quadrant (one-half of the upperjaw), measuring the skull’s greatestlength (from the posteriormost marginto the anteriormost portion, notincluding the incisors), and comparingthese lengths using the dental formulafor each species (table 22) and theskull key.


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 platesremain in the joints) and consist of two protuberances or a single protuberance that is larger and more tapered than in adults.

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

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Appendix

Locations of the six standardbody measurements—total length(TL), tail length (TV), foot length(HF), ear length (E), forearm length(FA), and tragus length (TR)—areillustrated in figures 45 and 46. Thesemeasurements can be taken from deadspecimens prior to preparation or fromlive specimens prior to release.Because we designed the keys usingcharacteristics and measurementstaken from adult individuals, theymay not be reliable when identifyingjuveniles. Juvenile and adult ageclasses can be determined byexamining the extent of epiphyseal-diaphyseal fusion in the finger joints(Anthony 1988). Cartilaginous platesmay not be apparent in the finger jointsof adults; the joints may consist of asingle, knobby protuberance (fig. 45).Cartilaginous plates are apparent inthe finger joints of juveniles; the jointsconsist of two protuberances with aslight taper between or a singleprotuberance that is much longerand more tapered at both ends thanthe protuberance in adult joints (fig.45). The most effective way to observethe cartilaginous plates is by back-lighting the wing and looking forsemitransparent sections in jointsof the phalanges. We also haveincluded a summary table listingthe characteristics of each species(table 23).


Tabl

e 23

—A

com

paris

on o

f cha

ract

eris

tics o

f bat

s of t

he S

avan

nah

Rive

r Site

Spec

ies

Pela

geFo

rear

mW

eigh

tFo

otEa

rTr

agus

shap

eaCa

lcara

Othe

r ide

ntify

ing

traits

mm

gm

m

Braz

ilian

free

-taile

d ba

t(Ta

darid

a br

asili

ensis

)Sh

ort, b

row

n to

dar

k gra

y43

11–1

48–

1119

–20

Tail f

ree f

rom

mem

bran

e

Rafin

esqu

e’s b

ig-e

ared

bat

(Cor

ynor

hinu

s raf

ines

quii)

Gray

or b

row

n ab

ove a

nd w

hitis

h be

low

417–

108–

1327

–37

Broa

d an

d lo

ngLa

rge e

ars

Easte

rn re

d ba

tBr

ight

red

or ru

st co

lor, f

emal

es w

ith fr

ostin

g,Sh

ort, b

lunt

,M

embr

ane b

etw

een

rear

legs

is(L

asiu

rus b

orea

lis)

mal

es w

ithou

t40

8–16

9–10

17–1

8an

d cu

rved

com

plet

ely f

urre

d

Hoar

y bat

Yello

wish

bro

wn

to d

ark b

row

n w

ith ve

ry h

eavy

Shor

t, blu

nt,

Very

larg

e; m

embr

ane b

etw

een

(Las

iuru

s cin

ereu

s)fro

sted

tips

5320

–35

6–13

17an

d cu

rved

rear

legs

is co

mpl

etel

y fur

red

Easte

rn p

ipist

relle

Blun

t and

stra

ight

Not

(Pip

istre

llus s

ubfla

vus)

Sand

y bro

wn

to o

rang

e on

back

—tri

colo

red

334–

78–

1013

–15

keel

edPi

nk fo

rear

ms,

blac

k win

gs

Silve

r-hai

red

bat

Blun

t and

Mem

bran

e bet

wee

n re

ar le

gs is

(Las

iony

cteris

noc

tivag

ans)

Dark

bla

ckish

bro

wn,

with

silve

r fro

sted

tips

417–

167–

1114

roun

ded

furre

d on

ante

rior h

alf

Big

brow

n ba

tBr

oad

and

Larg

e; pr

omin

ent g

land

s(E

ptes

icus f

uscu

s)Lo

ng b

row

n, w

ith tw

o-to

ned

blac

k bas

e47

13–2

510

–12

17–1

8ro

unde

dKe

eled

on m

usse

l

Even

ing

bat

Not

Stro

ng o

dor, p

rom

inen

t gla

nds

(Nyc

tece

ius h

umer

alis)

Dull b

row

n w

ith p

aler

bel

ly, yo

ung

muc

h da

rker

365–

147–

814

–15

Shor

t and

roun

ded

keel

edon

mus

sel, s

ingl

e upp

er in

cisor

Easte

rn sm

all-f

oote

d m

yotis

Shar

ply

Blac

k mas

k, re

lative

ly lo

ng ta

il(M

yotis

leib

ii)Bl

acki

sh b

row

n ab

ove,

pale

r bel

ow32

4–6

6–8

13–1

5Sh

arp

keel

ed(3

3 m

m)

Sout

heas

tern

myo

tisW

ooly,

dul

l, litt

le co

ntra

st b

etw

een

base

and

tips,

Slen

der a

ndNo

tGr

ay-b

lack

win

gs, lo

ng(M

yotis

austr

orip

ariu

s)ye

llow

ish to

gra

y abo

ve, w

hite

bel

ow40

5–12

10–1

313

–16

poin

ted

keel

ed to

e hai

rs

North

ern

long

-ear

ed m

yotis

Very

long

and

Larg

e pro

pata

gium

, ear

s(M

yotis

sept

entri

onal

is)No

t glo

ssy,

brow

n ab

ove,

gray

ish b

elow

355–

107–

1016

–19

slend

erKe

eled

exte

nd b

eyon

d no

se (3

mm

)

Little

bro

wn

bat

Long

glo

ssy t

an to

dar

k bro

wn

abov

e, gr

ay to

Slen

der a

ndNo

tLo

ng to

e hai

rs (e

xten

ding

bey

ond

(Myo

tis lu

cifug

us)

buff

belo

w38

7–8

8–11

13–1

6po

inte

dke

eled

claw

s), w

ings

cont

rast

ing

with

bod

y

a Bla

nk ce

lls in

dica

te ch

arac

teris

tics n

ot im

porta

nt fo

r spe

cies i

dent

ifica

tion.

65

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

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Appendix

Key to the Skins

1. a. One-third or more of the tailextending beyond uropatagium(fig. 47a)—Brazilian free-tailedbat (Tadarida brasiliensis)b. Tail not extending beyonduropatagium, or only slightly(fig. 47b)—2

2. a. Dorsal surface of uropatagiumat least partially furred—3b. Dorsal surface of uropatagiumnot furred or slightly furred atthe junction with the body—7

3. a. Pelage black, tips of hairsfrosted with white—4b. Pelage red, mahogany, oryellow—5

4. a. Total length > 120 mm,uropatagium heavily furredthroughout, ear white or yellowwith black rim—hoary bat(Lasiurus cinereus)b. Total length > 115 mm, posteriorone-third of uropatagium bare, earsolid black—silver-haired bat(Lasionycteris noctivagans)

5. a. White shoulder patch absent,yellow coloration, frostingabsent—northern yellow bat(Lasiurus intermedius)b. White shoulder patch present,red or mahogany coloration,frosting usually present (exceptmale Lasiurus borealis)—6

6. a. Pelage bright-brick red, tips ofhair frosted white (except males),face light red/yellow—eastern redbat (Lasiurus borealis)b. Pelage dark mahogany, tipsof hair frosted white, facemahogany/red—Seminole bat(Lasiurus seminolus)


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

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Illustration courtesy of B. Coslick

67

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

and southeastern myotis.

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Appendix

7. a. Ears > 25 mm long, distinctivepararhinal glands (large bumps,fig. 48) on either side of nose—Rafinesque’s big-eared bat(Corynorhinus rafinesquii)b. Ears < 25 mm long, pararhinalglands not distinct—8

8. a. Total length > 100 mm,forearm > 40 mm—big brownbat (Eptesicus fuscus)b. Total length < 100 mm,forearm < 40 mm—9

9. a. Tragus (projection withinthe ear) short, blunt, and curved(fig. 49a)—10b. Tragus long, pointed at tip,and straight (fig. 49b)—11

10. a. Dorsal fur tricolored whenparted; coloration black at base,yellowish brown in the middle anddark brown at tips; forearm pinkand < 32 mm—eastern pipistrelle(Pipistrellus subflavus)b. Dorsal fur dark brown, forearmdark and > 32 mm long—eveningbat (Nycticeius humeralis)

11. a. Ear > 16 mm long, extends > 2mm beyond the tip of nose whenlaid forward—northern long-earedmyotis (Myotis septentrionalis)b. Ear does not extend beyond thetip of nose when laid forward—12

12. a. Calcar keeled (figs. 50a and51a)—eastern small-footed myotis(Myotis leibii)b. Calcar not keeled (figs. 50band 51b)—13

13. a. Tips of hairs reddish; hair longand glossy—little brown bat(Myotis lucifugus)b. Tips of hairs not reddish; hairshort and wooly—southeasternmyotis (Myotis austroriparius)


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) theupper incisors of Brazilian free-tailed

bats, and (B) upper incisors of theother southeastern bats. Upper

incisors of the Brazilian free-tailed batconverge at the tips; upper incisors of

other southeastern bats do not.

Figure 53—Comparison of thelacrimal ridge protuberance of (A)eastern red and (B) Seminole bats.

Illustrations depict a dorsal surfaceview of the upper left skull quadrant.

Note that the two triangularstructures at the top are the upperincisor and the upper canine. Thehooked structure at the lower left

section is the anterior-most sectionof the zygomatic arch. The lacrimalridge protuberance (lacrimal shelf)typically is larger in (A) eastern red

bats than in (B) Seminole bats.

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Appendix

Key to Skulls

1. a. Upper incisors obviouslyconverge at tips, i.e., much closertogether at tips than at base (fig.52a—Brazilian free-tailed bat(Tadarida brasiliensis)b. Upper incisors wider at tip thanbase, equidistant at tip and base, ortips slightly converging (fig. 52b);premaxillary gap square—2

2. a. Nine teeth present in upperquadrant—Myotis spp.b. Fewer than nine teeth in upperquadrant—3

3. a. Eight teeth in upper quadrant—4b. Fewer than eight teeth in upperquadrant—6

4. a. Upper incisor bifid (two-cusped)—Rafinesque’s big-earedbat (Corynorhinus rafinesquii)b. Upper incisor unicuspid—5

5. a. Greatest length of skull> 13.5 mm; rostrum flatwith two concavities on dorsalsurface—silver-haired bat(Lasionycteris noctivagans)

b. Greatest length of skull< 13.5 mm, rostrum slopedwith no concavities on dorsalsurface—eastern pipistrelle(Pipistrellus subflavus)

6. a. Seven teeth in upper quadrant—7b. 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 > 15.5mm—hoary bat (Lasiurus cinereus)b. Greatest skull length < 15.5mm—9

9. a. Protuberance of the lacrimalridge (shelf) well developed(fig. 53a)—eastern red bat(Lasiurus borealis)b. Protuberance of the lacrimalridge poorly developed orabsent (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 poorlydeveloped, greatest length ofskull < 16 mm—evening bat(Nycticeius humeralis)

Menzel, Michael A.; Menzel, Jennifer M.; Kilgo, John C. [and others]. 2003. Batsof 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 (Pipistrellussubflavus), southeastern myotis (Myotis austroriparius), evening bat (Nycticeiushumeralis), 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 areextralimital capture records for two additional species: little brown bat (M. lucifigus) andnorthern yellow bat (Lasiurus intermedius). Acoustical sampling has documented thepresence 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. Thepresence of those two species, and a growing concern for the conservation of forest-dwellingbats, led to extensive and focused research on the Savannah River Site between 1996 and2002. Summarizing this and other bat research, we provide species accounts that discussmorphology and distribution, roosting and foraging behaviors, home range characteristics,habitat relations, and reproductive biology. We also present information on conservationneeds and rabies issues; and, finally, identification keys that may be useful wherever the batspecies we describe are found.

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

The Forest Service, United States Department ofAgriculture (USDA), is dedicated to the principleof multiple use management of the Nation’s forest

resources for sustained yields of wood, water, forage, wildlife, andrecreation. Through forestry research, cooperation with the Statesand private forest owners, and management of the National Forestsand National Grasslands, it strives—as directed by Congress—to provide increasingly greater service to a growing Nation.

The USDA prohibits discrimination in all its programs and activitieson the basis of race, color, national origin, sex, religion, politicalbeliefs, sexual orientation, or marital or familial status (Not allprohibited bases apply to all programs). Persons with disabilitieswho require alternative means for communication of programinformation (Braille, large print, audiotape, etc.) should contactthe USDA's TARGET Center at 202-720-2600 (voice and TDD).

To file a complaint of discrimination, write USDA, Director,Office of Civil Rights, Room 326-W, Whitten Building, 1400Independence Avenue, SW, Washington, DC 20250-9410 orcall 202-720-5964 (voice and TDD). USDA is an equalopportunity employer.

A publication of the Savannah River Site NationalEnvironmental Research Park Program. Publicationnumber: SRO-NERP-26

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