1

United States

Department of

Agriculture

Forest Service

Northeastern

Research Station

Research Note NE-378

Abstract

We attached a radio transmitter to an adult male Indiana

bat (Myotis sodalis) in June 2001 on the Fernow

Experimental Forest in the Allegheny Mountains of

north-central West Virginia. The bat was tracked for 4

successive days before the transmitter failed. The bat

roosted in three living trees over the study period. Two

roosts used for a single night each were in large shagbark

hickories (> 45 cm d.b.h.); the roost used for two

successive nights was located in a large sugar maple (69.1

cm d.b.h.). Roost trees were characterized by large areas of

exfoliating bark and all were canopy-dominant within

surrounding stands. One shagbark hickory was a residual

tree left following a patch clearcut 6 years earlier.

Although few inferences can be drawn from one Indiana

bat, many characteristics of this individuals’s roost

selections in the central Appalachians were consistent

with tree roosts observed in other regions during the nonhibernation period of this species.

The majority of the Indiana bat’s (Myotis sodalis) winter

hibernacula and summer maternity range is in the lower

Ohio Valley and Ozark Plateau of the Midwest (Menzel et

al. 2001b). However, there are 92 Priority II and III

hibernacula of this endangered species in the central and

southern Appalachians from northeastern Alabama to

central Pennsylvania (Humphrey 1978; USDI Fish and

Wildl. Serv. 1999; Menzel et al. 2001b). Summer

maternity activity of female Indiana bats is believed to be

rare in forest habitats in the central Appalachians of

Virginia and West Virginia (Brack et al. 2001; Owen et

al. 2001). However, most male Indiana bats remain in the

W. MARK FORD and JENNIFER M. MENZEL,

research wildlife biologists, USDA Forest Service,

Northeastern Research Station, Parsons, West Virginia;

MICHAEL A. MENZEL, graduate research assistant,

and JOHN W. EDWARDS, assistant professor, West

Virginia University, Division of Forestry, Morgantown,

West Virginia.

Summer Roost-Tree Selection

by a Male Indiana Bat on the

Fernow Experimental Forest

W. Mark Ford

Jennifer M. Menzel

Michael A. Menzel

John W. Edwards
2

hibernacula vicinity and use trees and snags as day roosts

during late spring, summer, and early fall. Accordingly,

protection of tree roosts and forested habitat around

Indiana bat hibernacula in the Appalachians is important

to safeguard males from direct mortality and/or harmful

modification of their roosting and foraging habitat.

Within the central and southern Appalachians, published

roost research on male Indiana bats has been limited to a

study of immediate post-hibernation emergence in the

Ridge and Valley of western Virginia (Hobson and

Holland 1995) and two pre-hibernation studies

conducted just before and during fall “swarm” on the

Cumberland Plateau in eastern Kentucky (Kiser and

Elliot 1996; MacGregor et al. 1999). Although critical for

natural resource managers in a region where forest

management is substantial on private lands (DiGiovanni

1990) while decreasing rapidly on public lands (Ford et

al. 2000), there are few data on roost characteristics of

male Indiana bats during the non-hibernation spring and

summer seasons. The objective of this study was to

provide preliminary data on male Indiana bat day roosts

in the central Appalachians of West Virginia and relate

those findings to current forest management practices.

Study Area and Methods

We examined summer day roosts of a male Indiana bat on

the Fernow Experimental Forest, a 1,900-ha research

forest located in Tucker County, West Virginia. Elevations

in this portion of the Allegheny Mountains subsection of

the Unglaciated Appalachian Mountains and Plateau

Physiographic Province generally range from 600 to 1300

m. Topography is characterized by steep side slopes with

broad ridge tops and narrow valleys (Fenneman 1938).

The climate is cool and moist with annual precipitation

exceeding 155 cm (Madarish et al. 2002). On upland

sites, mature (> 70 years), second-growth Allegheny/

northern hardwood forests are dominated by American

beech (Fagus grandifolia), sugar maple (Acer saccharum),

red maple (A. rubrum), black cherry (Prunus serotina),

northern red oak (Quercus rubra), yellow birch (Betula

alleghaniensis), black birch (B. lenta), Fraser magnolia

(Magnolia fraseri), and basswood (Tilia americana).

Riparian areas on the Fernow are dominated by eastern

hemlock (Tsuga canadensis) and rosebay rhododendron

(Rhododendron maximum). Because the Fernow is

designated as a forestry research area, it contains younger

forest stands (< 10 years) that originated from clear- and

deferment cutting and older stands altered by diameterlimit and selection cutting.

Portions of the Fernow and surrounding landscape in

eastern West Virginia are underlain by the Greenbrier

Limestone strata (Madarish et al. 2002) with karst

formations and numerous caves. Locally, several of these

caves serve as minor hibernacula for Indiana bats. Big

Springs Cave, located near the center of the Fernow,

annually winters approximately 200 male and female

Indiana bats. During the summer maternity season,

female Indiana bats probably are absent throughout most

of the Allegheny Mountains of West Virginia, including

the Fernow (Owen et al. 2001). Male Indiana bats that

hibernate in Big Springs Cave remain in and around the

Fernow (≤ 25 km radius) during the spring, summer and

fall where they day roost in trees (L. E. Thomasma,

USDA Forest Service, commun.). As a result, timber

harvests for research purposes on the Fernow are limited

to October 1 through April 30 to avoid “take” of an

endangered species through habitat modification or direct

mortality that could occur by felling an occupied roost

tree.1

On 16 June, 2000, at approximately 2200 hr, we captured

a male Indiana bat with a single mist net placed over

Elklick Run near the center of the Fernow Experimental

Forest. We confirmed species identification by the

presence of a keeled calcar and short toe hairs that did not

extend beyond the knuckle or claw (Whitaker and

Hamilton 1998). The bat had the following

measurements: forearm 38 mm, ear 8.5 mm, tragus 6

mm, and mass 7.0 g. We assigned the bat to the adult age

class by examining the degree of epiphyseal-diaphyseal

fusion (Anthony 1988; Racey 1988). We attached a 0.51-

g model LB-2 radio transmitter (Holohil Systems Ltd.,

Woodlawn, ON)2

 to the hair between the bat’s scapula

using Skin Bond® surgical adhesive (Pfizer Hospital

Products Group, Largo, FL). Transmitter mass was

approximately 7 percent of the bat’s body weight, slightly

in excess of the recommended 5 percent of body mass

(Aldridge and Brigham 1988). We used LA12-Q AVM

receivers (AVM Instruments, Colfax, CA) and folding,

three-element Yagi antennas to locate the Indiana bat’s

day roosts. The bat was tracked for 4 successive days

before the transmitter was shed.

At each day roost or within a 10-m-radius circular plot

with the roost as plot center, we recorded roost-tree

species, d.b.h. (cm), height (m), snag class, bark-cover

1

Adams, M. B., Knibbs, J., Rodrigue, J. L., Edwards, P. J.,

Wood, F., Ford, W. M., Kochenderfer, J. A., Schuler, T. M.,

Crews, J. 2000. Fernow Experimental Forest: environmental

impact statement, Final Rep. On file at the Northeastern

Research Station, Newtown Square, Pennsylvania.

2

The use of trade, firm, or corporate names in this report is for

the information of the reader. Such use does not constitute an

official endorsement or approval by the USDA Forest Service of

any product or service to the exclusion of others that may be

suitable.
3

class, site aspect, elevation (m), surrounding forest-stand

community, canopy cover, midstory and woody-seedling

density, litter depth, and distance (m) to nearest water, to

Elklick Run, to nearest road, to the capture site, and to

Big Springs Cave. We used a laser rangefinder to

determine roost height and the distance to the nearest

road. Site aspect was linearized using (1-cosine[aspect

degrees] + 1-sine[aspect degrees]) with values increasing

from mesic, northern aspects to xeric, southwestern

aspects (Odom et al. 2001). Following Maser et al.

(1979), we assessed snags from 1-7, with 1 a live tree and

7 a decomposing broken bole. We rated bark-cover class

as I (none), II (< 10 percent), III (10-25 percent) and IV

(> 25 percent). We grouped visual estimates of canopy

cover, midstory density, and woody-seedling and

herbaceous groundlayer density in five classes: I (0-5

percent), II (5-25 percent), III (26-50 percent), IV (51-75

percent), V (76-95 percent), and VI (96-100 percent). We

rated litter depth as I (none), II (≤ 2.54 cm), III (2.55-6

cm), and IV (> 6 cm). We calculated elevation and

distances from roosts to nearest water, Elklick Run,

capture location, Big Springs Cave, and other roost trees

using ArcView 3.2 GIS (ERSI, Redlands, CA).

Results and Discussion

Over a 4-day period, we tracked the male Indiana bat to

three different roosts (Fig.1), all of which were Stage 1

live trees with abundant exfoliating bark (> 25 percent

bole surface area) and no visible cavity or wound (Table

1). Roosts 1 and 2 were large shagbark hickories (Carya

ovata) used by the bat for 1 day each (Table 1). Roost 3

was a large sugar maple that was used by the bat for 2

successive days (Table 1). All roosts were located in mesic

Allegheny/northern hardwoods communities on sites with

northwest to northeast aspects (Table 1). Roosts 1 and 3

were in mature forest stands characterized by numerous

large trees. Roost 2 was a residual tree in a patch clearcut.

The stand had been harvested in 1995 and was

dominated by abundant woody regeneration 3 to 4 m

high. Roosts 1 and 3 were canopy-dominant trees, though

neither extended above the canopy. Each day roost was <

50 m from a road, < 300 m to the nearest water, < 1,000

m to Elklick Run and the capture site, and < 2,000 m to

Big Springs Cave (Table 1). Distances from Roost 1 to

Roost 2, Roost 2 to Roost 3, and Roost 1 to Roost 3 were

340, 477, and 705 m, respectively.

Table 1.—Summer day-roost characteristics for an adult male Indiana bat on the Fernow

Experimental Forest, Tucker County

Variable Roost 1 Roost 2 Roost 3 Mean SE

Tree species C. ovata C. ovata A. saccharum

d.b.h. (cm) 45.5 68.0 69.1 60.9 7.7

Height (m) 32.5 30.0 25.8 29.4 2.0

Snag class 1 I I

Bark cover IV IV IV

Linearized aspect 2.7 3.5 3.9 3.4 0.4

Slope (%) 38 57 83 59.3 13.1

Elevation (m) 800 800 702 767.2 32.9

Stand type N. Hardwood N. Hardwood N. Hardwood

Canopy cover III I V

Midstory layer IV V III

Seedling layer I II II

Litter depth II II III

Distance (m) to:

Nearest water 243 85 53 127.2 58.7

Elklick Run 740 803 618 720.3 54.4

Nearest road 43 49 13 35.0 11.2

Capture site 868 905 643 805.3 82.0

Big Springs Cave 1275 1588 1980 1614.3 204.2
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Tree-roosting bats generally select trees that are larger in

diameter and taller than the surrounding stand (Sasse and

Perkins 1996; Vonhof and Barclay 1996; Callahan et al.

1997; Foster and Kurta 1999). Although both male and

female Indiana bats show large rangewide variation in the

size (8 to 86.6 cm d.b.h) and types of trees or snags

chosen for roosts (Menzel et al. 2001b), Kurta et al.

(1996) found that roost trees tended to be larger on

average than other trees in the surrounding stand. The

three roosts on the Fernow were similar to or larger than

the majority of live trees or dead snags used by females at

maternity roosts in the Midwest (Gardner et al. 1991;

Kurta et al. 1996; Callahan et al. 1997) or as day roosts

for males in the early spring or late fall in the

Appalachians (Hobson and Holland 1995; Kiser and

Elliot 1996; MacGregor et al. 1999). Female bats often

choose a large tree or snag that extends to the canopy or

above to take advantage of increased solar exposure that

keeps the roost warm and aids the growth and

development of juvenile bats (Menzel et al 2001a).

Presumably, male Indiana bats roosting in and around Big

Springs Cave are not so constrained because all three

roosts were situated on more northern (cooler) aspects.

Callahan et al. (1997) speculated that male Indiana bats

seek cooler roosts to conserve energy. Other measured

variables such as canopy cover were similar to or within

ranges reported for tree roosts from throughout the

Indiana bat’s distribution (Menzel et al. 2001b).

Rangewide, Indiana bats show an affinity for roost trees

with exfoliating bark (Rommé et al. 1995; Callahan et al.

1997). Menzel et al. (2001b) identified snags of 23 tree

1 km

Big Springs Cave

Capture Location

Roost 3

Roost 2

Roost 1

Elklick Run

Figure 1.—Capture site and day roosts of a male Indiana bat in relation to Big Springs Cave

and Elklick Run on the Fernow Experimental Forest in West Virginia.
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species used by Indiana bat as day roosts. Shagbark

hickory, noted for extensive amounts of exfoliating bark,

was the lone live species often used for roosts (Humphrey

et al. 1977; Gardner et al. 1991; Callahan et al. 1997).

The large sugar maple identified as a roost also had a

substantial amount of exfoliating bark. Because shagbark

hickory is such a minor forest component within the

Fernow’s Allegheny/northern hardwood type, male

Indiana bats might be showing strong preference for this

tree species. Unfortunately, in the absence of even-age

silviculture and continued fire suppression, heavy seeded,

shade-intolerant species such as shagbark hickory

continue to decrease in importance on the Fernow and

surrounding landscape (Schuler and Gillespie 2000).

However, the large increase in dominance of sugar maple

across the Fernow partially could offset losses of shagbark

hickory.

Regardless, it seems appropriate to retain large live trees

or dead snags within predetermined radii around Indiana

bat hibernacula in the central and southern Appalachians

where males may roost in spring through early fall, and to

prohibit forest harvesting within that area during the

non-hibernation season (Kiser and Elliot 1996). Within

these hibernacula zones and at the appropriate season,

two-age or deferment harvests (Miller et al. 1995;

Johnson et al. 1998) that retain large shagbark hickory,

sugar maple, or other trees with exfoliating bark would

provide timber products and adequate regeneration while

protecting the Indiana bat and its habitat.

Spatially, the locations of the three day roosts relative to

where the male Indiana bat was captured on its presumed

foraging area along Elklick Run are well within the ranges

of > 200 m to 2 km from day roosts to foraging areas

reported in the literature (Humphrey et al. 1977; Kurta et

al. 1993; Menzel et al. 2001b). On an Appalachian

landscape somewhat comparable to the Fernow, Hobson

and Holland (1995) observed a 1-km distance between

male Indiana bat day roosts and riparian-zone foraging

areas in western Virginia during the post-hibernation

period in spring. Acoustical survey sampling has indicated

that Indiana bats on the Fernow forage extensively along

Elklick Run during the summer.3,4 Distances between

observed roost trees also were within ranges reported for

day roosts in the early fall on the Cumberland Plateau

(MacGregor et al. 1999). Our observation that the male

Indiana bat we tracked used three day roosts over 4 days

was consistent with frequent roost switching (1 to 2 days)

observed for male Indiana bats in the pre-swarm period in

the fall on the Cumberland Plateau (Kiser and Elliot

1996; MacGregor et al. 1999). Additional research and

monitoring is needed to determine whether the

midsummer roost characteristics we observed are

representative of male Indiana bats on the Fernow and/or

similar to pre- and post-hibernation data collected from

other portions of the central and southern Appalachians.

Acknowledgments

Field assistance was provided by Jane Rodrigue and Kate

Leonard. Funding for this work was provided by the

Northeastern Research Station Scientist and the West

Virginia University, Swiger Graduate Fellowship.

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