United States

Department of 

Agriculture

Forest Service

Northern 

Research Station

Research Paper NRS-8

Notes on Foraging 

Activity of Female 

Myotis leibii in Maryland

Joshua B. Johnson

J. Edward Gates

W. Mark Ford
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Abstract

Information on home range and habitat characteristics of eastern small-footed myotis (Myotis 

leibii) consist only of anecdotal accounts and unpublished research despite the need for such 

data for conservation of this rare species. We used radio telemetry to determine foraging site 

selection of four female eastern small-footed myotis in Allegany County, Maryland, in spring 

2007. These bats foraged within 1.8 km of their diurnal roosts and had home ranges of <100 

ha. Distance-based analysis of habitat use for one foraging bat showed that it foraged farther 

from the Potomac River and adjacent wetlands, and closer to hilltop forests than random, 

conforming to expectations based on their low wing loading and broadband, high frequency 

echolocation call characteristics.

Cover Photos

Small-footed myotis, by Craig Stihler, West Virginia Division of Natural Resources, 

used with permission.
1

Th e Authors

JOSHUA B. JOHNSON is a graduate 

research assistant, Division of Forestry and 

Natural Resources, West Virginia University, 

Morgantown, West Virginia 26506.

J. EDWARD GATES is an associate 

professor, University of Maryland Center 

for Environmental Science, Appalachian 

Laboratory, Frostburg, Maryland 21532.

W. MARK FORD formerly was a research 

wildlife biologist, U.S. Forest Service, 

Northern Research Station, Parsons, WV 

26287. He currently is a research wildlife 

biologist, U.S. Army Engineer Research and 

Development Center Environmental Lab, 

Vicksburg, Mississippi 39180.

INTRODUCTION

Th e eastern small-footed myotis (Myotis leibii) is considered 

among the rarest bat species in North America. Consequently, 

little is known about its natural history and ecology compared 

to sympatric bat species (Best and Jennings 1997). Foraging 

areas undoubtedly are a critical component of eastern smallfooted myotis habitat, as with any bat species, however, 

information on foraging habitat selection and home range 

size currently is restricted to anecdotal observations and 

unpublished research (Erdle and Hobson 2001).

Eco-morphology predicts that eastern small-footed myotis 

foraging characteristics should be similar to other bat species 

that have comparable wing loading and echolocation call 

structure (Patriquin and Barclay 2003, Ford et al. 2005). For 

example, bats that have low wing loading and broadband, 

high frequency echolocation call characteristics are capable 

of effi ciently foraging in cluttered areas such as forest canopy 

(Norberg and Raynor 1987, Kalcounis and Brigham 1995). 

Eastern small-footed myotis are known to emit broadband, 

high frequency echolocation calls, which are consistent with 

foraging in structurally cluttered forested areas (Mukhida 

et al. 2004). In eastern West Virginia, recent unpublished 

work has shown that radio-tagged eastern small-footed 

myotis foraged mostly in forested areas.1

 Research on food 

habits of eastern small-footed myotis suggests they use a 

gleaning strategy, possibly conducive to foraging in forest 

canopies (Johnson and Gates 2007, Moosman et al. 2007). 

In this study, our objectives were to 1) determine foraging 

characteristics, i.e., home range size and foraging habitat 

selection, of eastern small-footed myotis; and 2) consider the 

conformity of their foraging characteristics to predictions 

based on their wing morphology, echolocation call 

characteristics, and food habits.

1

C. Stihler, West Virginia Division of Natural Resources, 

personal communication.
2

SITE DESCRIPTION

We conducted our research within the Chesapeake 

and Ohio Canal National Historical Park (CHOH) 

in Allegany County, Maryland within the Ridge and 

Valley physiographic province. Our eff orts were focused 

at an abandoned railroad tunnel used by this and other 

species of bats as a hibernaculum (Johnson and Gates 

2008). Th e area is characterized by steep slopes, shale 

barrens, and rock outcrops overlooking the Potomac 

River. Forests on the slopes and ridges are predominately 

oak-hickory (Quercus and Carya spp.) with some 

scattered Virginia pine (Pinus virginiana) and red cedar 

(Juniperus virginiana). Pastures and row crops comprise 

a minor component of the cover types on the hilltops. 

Yellow poplar (Liriodendron tulipifera), sycamore 

(Platanus occidentalis), silver maple (Acer saccharinum), 

red maple (A. rubrum), and river birch (Betula nigra) 

occur along the Potomac River banks and the adjacent 

CHOH. Ephemeral standing water and wetlands occur 

along some sections of the canal. An active railroad is 

located on the West Virginia side of the Potomac River. 

Elevation in the immediate area ranges from about 130 

m on the Potomac River to about 275 m on the hilltops.

METHODS

We used harp traps (1.8 m  2.3 m; Bat Conservation 

and Management, Carlisle, PA) to capture bats at the 

tunnel as they emerged from hibernation during spring 

2007. We placed two harp traps side by side in both the 

east and west tunnel entrances to capture bats entering 

or exiting the tunnel. Th e harp traps were surrounded 

with tarpaulin and/or plastic netting to prevent bats from 

bypassing the traps. Sampling2

 was conducted 3 nights/

week following sunset for four hours for 31 nights from 

12 March through 16 May 2007. Each captured bat was 

identifi ed to species. We measured mass (g) and forearm 

length (mm), and the sex of each bat was determined 

before its release (Kunz 1988, Menzel et al. 2002). We 

traced the shape of a fully extended wing of a random 

sample of eastern small-footed myotis to determine 

wing morphology and wing loading for predictions 

about fl ight performance and habitat use (Kalcounis 

and Brigham 1995). Wing aspect ratio was calculated 

by dividing the wing length by fi fth phalanx length. 

We used a polar planimeter (Los Angeles Scientifi c 

Instrument Co., Inc., Los Angeles, CA) to measure wing 

surface area of each tracing. Wing loading was calculated 

by dividing body mass by the combined surface area 

of both wings and uropatagium, following Farney and 

Fleharty (1969).

Radio telemetry allowed us to examine foraging activity 

of female eastern small-footed myotis. We used surgical 

cement (Torbot Group, Cranston, RI) to affi x a 0.35-g 

radio transmitter (Model LB-2N; Holohil Systems 

Ltd., Carp, Ontario, Canada) between the scapulae of 

captured female eastern small-footed myotis. Th e ratio of 

transmitter mass to body mass (mean = 8.0 percent, std. 

dev. = 0.4 percent, range = 7.8-8.8 percent) was similar 

to ratios reported in other studies on bats, including the 

similarly-sized eastern pipistrelle (Perimyotis subfl avus; 

Best and Jennings 1997, Carter et al. 1999, Perry and 

Th ill 2007, Veilleux et al. 2003).

We used radio receivers and three-element Yagi 

antennaes (Advanced Telemetry Systems, Inc., Isanti, 

MN) to simultaneously obtain two directional bearings 

from known locations (i.e., telemetry stations) at 

5-minute intervals (Menzel et al. 2005). We positioned 

telemetry stations to minimize distances to radiotagged bats and to reduce associated location error. 

Telemetry station locations and bearings were entered 

into Locate III (Pacer Computing, Inc., Tatamagouche, 

NS, Canada) to obtain Universal Transverse Mercator 

(UTM) coordinates of each foraging location (Nams 

2006). We entered coordinates of foraging locations and 

diurnal roost locations for each bat into ArcView 3.2 

(Environmental Systems Research Institute, Redlands, 

CA) and used the Animal Movement Extension to 

calculate a home range, regardless of minimum number 

of locations, for each bat using the fi xed kernel method 

based on a 95 percent confi dence interval to exclude 

outliers (Hooge and Eichenlaub 1997, Seaman et al. 

1999, Worton 1989).

2

Bat capture and handling protocols were approved by the 

Institutional Animal Care and Use Committee (ACUC) of 

the University of Maryland Center for Environmental Science 

(Protocol Number F-AL-05-06) and followed the guidelines 

of the American Society of Mammalogists (1998).
3

We examined foraging habitat selection of bats for 

which we obtained >30 locations (Aebischer et al. 1993). 

A raster land use/land cover (LULC) theme from the 

United States Geological Survey (USGS) Gap Analysis 

Program was converted to a LULC vector shapefi le in 

ArcView 3.2 (USGS 2000). From the LULC vector 

shapefi le, we created a separate vector shapefi le for each 

LULC type, including pasture/hayfi elds, wetlands, 

deciduous forest, coniferous forest, mixed forest, and all 

forests combined. Separate vector shapefi les, including the 

Potomac River, railways, and paved roads, were included 

in the analysis for nine LULC types. We used distancebased analysis to examine habitat use of radio-tagged bats 

because it reduces the eff ect of radio telemetry error and 

Type 1 error commonly associated with other habitat 

use analyses, including compositional analysis (Bingham 

and Brennan 2004, Conner et al. 2002). Distance-based 

analysis compares the distances of foraging locations to 

each LULC type with the distance of random locations to 

the same LULC type (Conner et al. 2002). For each bat, 

we determined the maximum distance it foraged from its 

diurnal roost (Johnson and Gates 2008). Th is distance 

served as a radius for a buff er from which random 

locations were generated; this buff er was centered on a 

bat’s diurnal roost location. To determine if eastern smallfooted myotis were foraging randomly among LULC 

types, we used ArcView 3.2 to pair each foraging location 

with a random location within the buff er. Minimum 

Euclidean distances from every foraging and random 

location to every LULC type were determined. We used 

a Mann-Whitney test to examine diff erences between 

foraging and random location distances to each LULC 

type (SAS Institute, Inc. 2004; PROC NPAR1WAY). 

Statistical signifi cance was set at P ≤ 0.05.

RESULTS

We captured 47 eastern small-footed myotis: 33 

males, 13 females, and one escaped before sex could 

be determined. We traced the wings of seven eastern 

small-footed myotis. Mean single wing length (± 1 SE, 

range) was 86.57 mm (± 4.17, 63.0-96.0), and mean 

fi fth phalanx length was 44.29 mm (± 0.75, 40.0-46.0). 

Mean wing aspect ratio was 1.95 (± 0.10, 1.40-2.18). 

Mean total wing surface area, not including uropatagium, 

was 51.75 cm2

 (± 3.42, 34.55-60.83). Mean mass of 

bats from which wing tracings were obtained was 4.39 g 

(± 0.17, 3.5-5.0). Mean wing loading was 0.065 g/cm2

(± 0.002, 0.058-0.071).

We conducted radio telemetry on four female eastern 

small-footed myotis from 13 March through 4 April 

2007. Radio contact was maintained with all but one 

bat until the transmitters died or detached from the bats 

(mean = 8 days, range = 7-9 days). We lost the signal of 

one bat (frequency 151.398) after its seventh night of 

activity when it crossed the Potomac River into West 

Virginia. Radio-tagged bats were tracked 2-5 nights and 

we obtained 5-74 foraging locations from each (Table 1). 

Bats foraged within 1.8 km of their diurnal roosts and 

each had minimum home range of <100 ha (Table 1).

We conducted distance-based analysis of habitat use on a 

single bat (frequency 151.098), located 74 times during 

5 nights between 27 March and 4 April 2007 (Figure 

1). Compared to random locations, this bat foraged 

farther from railroads, the Potomac River, and wetlands. 

Conversely, this bat foraged closer to paved roads, 

pastures, coniferous forest, and mixed forest than random 

Table 1.—Foraging characteristics of female Myotis leibii in Allegany County, Maryland, March-April 2007

Foraging

Distance (m) b

Radio 

frequency

Number of 

nights

Number of 

locations

Home range size 

(ha) a Mean SE Range

151.098 5 74 99.7 817.4 38.0 68.7 - 1728.0

151.379 2 5 10.2 189.5 64.0 18.4 - 314.8

151.398 3 14 62.7 355.6 98.7 16.4 - 1203.4

151.417 2 10 42.4 897.8 31.9 660.1 - 1009.8

a

 95 percent fi xed kernel. b

 Distance from foraging location to diurnal roost.
4

Figure 1.—Fixed kernel home range estimates for a female eastern small-footed myotis (Myotis leibii) 

in Allegany County, Maryland, spring 2007. 
5

locations. Distances from deciduous forest and all forest 

types combined did not diff er (P > 0.352) between 

foraging locations and random locations (Table 2). Th e 

remaining three bats that we tracked foraged over the 

Potomac River, in adjacent riparian forests, and on the 

forested hilltops.

DISCUSSION

Our results provide insight into the home range and 

foraging habitat of eastern small-footed myotis, and are 

consistent with anecdotal observations and results of 

unpublished research. Th e foraging activity characteristics 

of eastern small-footed myotis may be similar to that 

of other myotine bats in the region, particularly that of 

the northern myotis. Eastern small-footed myotis have 

low wing loading and an echolocation call structure that 

is similar to that of northern myotis, which also have 

similar home ranges and diet preferences (Brack and 

Whitaker 2001, Farney and Fleharty 1969, Johnson and 

Gates 2007, Moosman et al. 2007, Mukhida et al. 2004, 

Owen et al. 2003). Bat species with low wing loading 

and high, frequency-modulated echolocation calls are 

capable of foraging effi ciently in cluttered forest interiors 

and commonly have smaller home ranges than sympatric 

species in the mid-Atlantic and central Appalachian 

region (Kalcounis and Brigham 1995, Owen et al. 2003, 

Patriquin and Barclay 2003, Wund 2006). However, 

because we obtained >30 foraging locations for only a 

single eastern small-footed myotis, our results should 

be interpreted with caution as this individual may not 

be representative of the species as a whole (Girard et al. 

2006). Also, we exceeded the recommended 5 percent 

“rule” for radio transmitter burden for the four bats 

that we tracked (Aldridge and Brigham 1988) and we 

obtained relatively few (≤14) foraging locations for three 

of those. Consequently, our home range calculations 

should be interpreted as coarse estimates for the species.

Caveats aside, our results showed that a single eastern 

small-footed myotis foraged more in forested areas (93.6 

percent of foraging locations), particularly deciduous 

forests (62.8 percent), than in open areas such as 

pastures (2.6 percent). Also, we observed that many 

foraging locations were along ridge tops near day roosts, 

which were located in rock outcrops in shale barren 

slopes overlooking the Potomac River (Johnson and 

Gates 2008). Th e full extent of eastern small-footed 

myotis occupancy of rock outcrops in the Appalachian 

Mountains remains to be determined (Johnson and 

Gates 2008), but other known sites in Maryland have 

included rock outcrops such as those at High Rock, Big 

Savage Mountain nearby to the west.3

 Similar to other 

wildlife species in the region, particularly the Allegheny 

woodrat (Neotoma magister), eastern small-footed myotis 

are dependent on emergent rock habitats, as well as 

the immediately surrounding forests where they forage 

(Castleberry et al. 2001, Ford et al. 2006). How ongoing 

Table 2.—Distance-based habitat use analysis between female Myotis leibii (n = 1) foraging 

locations (n = 74) and random locations (n = 74) in Allegany County, Maryland, March-April 2007

Foraging location Random location

Variable Mean a SE Mean SE P

Railroad 762.4 46.4 384.3 45.4 <0.001

Potomac River 760.2 47.2 383.8 39.6 <0.001

Wetlands 585.9 24.1 365.2 22.8 <0.001

Paved roads 310.0 41.9 604.9 46.1 <0.001

Pasture 97.1 12.2 140.1 14.1 0.018

Coniferous forest 69.0 8.3 94.0 8.8 0.023

Mixed forest 39.4 5.4 54.7 5.6 0.017

Deciduous forest 6.5 1.4 6.2 1.5 0.352

All forest combined 1.0 0.4 1.4 0.6 0.958

a

 Distances (m). Tests were performed on ranked data, but actual values are shown.

3D. Feller, Maryland Department of Natural Resources, 

personal communication.
6

changes to ridgeline forests in the region for activities 

such as wind energy development (Kunz et al. 2007, 

Arnett et al. 2008, Baerwald et al. 2008) or recreational 

development (Ford et al. 2006) impacts eastern smallfooted myotis foraging habitat are unknown. Accordingly 

a better understanding of the required spatial extent 

and structure of forest cover along ridgelines and 

rock outcrops, as well as additional foraging activity 

requirements, is needed to aid conservation eff orts for 

this rare species (Johnson and Gates 2008).

ACKNOWLEDGMENTS

We thank S. Carr, K. Lott, J. MacDougall, and J. 

Saville for assisting us in the fi eld during our survey 

eff orts. J. Churchill and K. Lott provided assistance 

with Geographic Information Systems. Th e Maryland 

Department of Natural Resources, Engineering and 

Construction Division, Land and Water Conservation 

and Power Plant Research Program provided funding. 

Th e comments of R. Perry, D. Feller, and J. Stanovick 

greatly improved this manuscript.

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Johnson, J.B.; Gates, J.E.; Ford, W.M. 2009. Notes on foraging activity of female 

Myotis leibii in Maryland. Res. Pap. NRS-8. Newtown Square, PA: U.S. 

Department of Agriculture, Forest Service, Northern Research Station. 8 p.

Information on home range and habitat characteristics of eastern small-footed 

myotis (Myotis leibii) consist only of anecdotal accounts and unpublished research 

despite the need for such data for conservation of this rare species. We used radio 

telemetry to determine foraging site selection of four female eastern small-footed 

myotis in Allegany County, Maryland, in spring 2007. These bats foraged within 

1.8 km of their diurnal roosts and had home ranges of <100 ha. Distance-based 

analysis of habitat use for one foraging bat showed that it foraged farther from the 

Potomac River and adjacent wetlands, and closer to hilltop forests than random, 

conforming to expectations based on their low wing loading and broadband, high 

frequency echolocation call characteristics.

KEY WORDS: eastern small-footed myotis, habitat use, home range, Maryland, 

Myotis leibii
Capitalizing on the strengths of existing science 

capacity in the Northeast and Midwest to attain a more 

integrated, cohesive, landscape-scale research program

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