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United States
Department of

Forest Service

Summer Roost-Tree Selection
by a Male Indiana Bat on the

Norarensrerh Fernow Experimental Forest
Research Station
Research Note NE-378 W. Ma rk Ford
Jennifer M. Menzel
Michael A. Menzel
John W. Edwards
oF
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The majority of the Indiana bat’s (Myotis sodalis) winter
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 non-
hibernation period of this species.

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.

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 (7suga 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 diameter-
limit 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

i)

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 (S 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 | 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.!

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)’ 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

'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.

°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.

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

Experimental Forest, Tucker County

Roost 3 Mean SE

Variable Roost 1 Roost 2
‘Tree species C. ovata C. ovata
d.b.h. (cm) 45.5 68.0
Height (m) 32.5 30.0
Snag class l I
Bark cover IV IV
Linearized aspect 2 ae
Slope (%) 38 pi
Elevation (m) 800 800
Stand type N. Hardwood N. Hardwood
Canopy cover Ul I
Midstory layer IV Vv
Seedling layer I Il
Litter depth I Ul
Distance (m) to:
Nearest water 243 85
Elklick Run 740 803
Nearest road 43 49
Capture site 868 905

A. saccharum

69.1 60.9 7.7
25.8 29.4 2.0

I

IV

59 3.4 0.4
83 Bs Be: 13.1
702 707.2 32.9

N. Hardwood

V

Ul

I]

Il

pe, L272 30.7
618 720.3 54.4
13 35.0 12
643 805.3 82.0
1980 1614.3 204.2

Big Springs Cave 1275 1588

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), II (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.

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.

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

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 with 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.** 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

3Owen, S. E, Menzel, M. A.; Ford, W. M.; Edwards, J. W.;
Menzel, J. M.; Chapman, B. R.; Wood, P. B.; Miller, K. V. Bat
activity in managed and unmanaged forest and riparian zones in
the Allegheny Mountains. In preparation.

*W. M. Ford, unpublished data on file at Northeastern Research
Station, Parsons, West Virginia.

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. Cover
photo courtesy of Timothy Carter.

Literature Cited

Aldridge, H. D.; Brigham, R. M. 1988. Load carrying
and maneuverability in an insectivorous bat: a test
of the 5% “rule” of radio-telemetry. Journal of
Mammalogy. 69: 379-382.

Anthony, E. L. 1988. Age determination. In: Kunz, T.
H., ed. Ecological and behavioral methods for the
study of bats. Washington, DC: Smithsonian
Institution Press: 47-57.

Brack, V.; Stihler, C. W.; Reynolds, R. J.; Butchkoski, C.
2001. Distribution, abundance, and reproduction
by the Indiana bat (Myotis sodalis) in the
Appalachian Mountains of the eastern U.S.: affect
of elevation. In: Kurta, A. ed. The Indiana bat,
biology and management of an endangered species;
2001 March 29-April 1; Lexington, KY. University of
Kentucky: 9. Abstract.

Callahan, E. V.; Drobney, R. D.; Clawson, R. L. 1997.
Selection of summer roosting sites by Indiana bats
(Myotis sodalis) in Missouri. Journal of Mammalogy.
78: 818-825.

DiGiovanni, D. M. 1990. Forest statistics for West
Virginia — 1975 and 1989. Resour. Bull. NE-114.
Radnor, PA: U. S. Department of Agriculture, Forest

Service, Northeastern Forest Experiment Station. 172

p-

Fenneman, N. M. 1938. Physiography of the eastern
United States. New York: McGraw-Hill. 714 p.

Foster, R. W.; Kurta, A. 1999. Roosting ecology of the
northern bat (Myotis septentrionalis) and

comparisons with the endangered Indiana bat
(Myotis sodalis). Journal of Mammalogy. 80: 659-672.

Gardner, J. E.; Garner, J. D.; Hofmann, J. E. 1991.
Summer roost selection and roosting behavior of
Myotis sodalis (Indiana bat) in Illinois. Final Rep.
Champaign, IL: Illinois Natural History Survey and

Illinois Department of Conservation. 56 p.

Hobson, C. S.; Holland, J. N. 1995. Post-hibernation
movement and foraging habitat of a male Indiana
bat, Myotis sodalis (Chiroptera: Vespertilionidae),
in western Virginia. Brimleyana. 23: 95-101.

Humphrey, S. R. 1978. Status, winter habitat, and
management of the endangered Indiana bat, Myotis
sodalis. Florida Science. 41: 65-76.

Humphrey, S. R.; Richter, A. R.; Cope, J. B. 1977.
Summer habitat and ecology of the endangered
Indiana bat, Myotis sodalis. Journal of Mammalogy.
58: 334-346.

Johnson, J. E.; Miller, G. W.; Baumbras, J. E.; West, C.
D. 1998. Assessment of residual stand quality and
regneration following shelterwood cutting in
central Appalachian hardwoods. Northern Journal of
Applied Forestry. 15: 203-210.

Kiser, J. D.; Elliot, C. L. 1996. Foraging habitat, food
habits, and roost tree characteristics of the Indiana
bat (Myotis sodalis) during autumn in Jackson
County, Kentucky. Final Rep. E-2. Frankfort, KY:
Kentucky Department of Fish and Wildlife Resources.
65 p.

Kurta, A.; Kath, J.; Smith, E R.; Orick, M. W.; Ross, R.
1993. A maternity roost of the endangered Indiana
bat (Myotis sodalis) in an unshaded, hollow,
sycamore tree (Platanus occidentalis). American
Midland Naturalist. 130: 405-407.

Kurta, A.; Williams, K. J.; Mies, R. 1996. Ecological,
behavioral, and thermal observations of a
peripheral population of Indiana bats (Myotis
sodalis). In: Barclay, R. M. R.; Brigham, R. M., eds.
Bats and forests symposium; 1995 October 19-21;
Victoria, BC. Victoria, BC: British Columbia Ministry
of Forests: 102-117.

MacGregor, J; Kiser, J. D.; Gumbert, M. W.; Reed, T. O.;
1999. Autumn roosting habitat of male Indiana
bats (Myotis sodalis) in a managed forest setting in
Kentucky. In: Stringer, J. W.; Loftis, D. L. eds.

6

Proceedings 12" central hardwood forest conference;
1999 February 28-March 1-2; Lexington, KY. Gen.
Tech. Rep. SRS-24. Asheville, NC: U.S. Department
of Agriculture, Forest Service, Southern Research
Station: 169-170.

Madarish, D. M.; Rodrigue, J. L.; Adams, M. B. 2002.
Vascular flora and macroscopic fauna on the
Fernow Experimental Forest. Gen. Tech. Rep. NE-
291. Newtown Square, PA: U.S. Department of
Agriculture, Forest Service, Northeastern Research
Station. 37 p.

Maser, C.; Anderson, R. G.; Cromak, K.; Williams, J. T.;
Martin, R. E. 1979. Dead and downed woody
material. In: Thomas, J. W., ed. Wildlife habitats in
managed forests—the Blue Mountains of Oregon and
Washington. Agric. Handb. 533. Washington, DC:
U.S. Department of Agriculture: 78-95.

Menzel, M. A.; Carter, T. C.; Ford, W. M.; Chapman, B.
R. 2001a. Tree-roost characteristics of subadult and
female adult evening bats (Nycticeius humeralis) in
the upper Coastal Plain of South Carolina.
American Midland Naturalist. 145: 112-119.

Menzel, M. A.; Menzel, J. M.; Carter, T. C.; Ford, W. M.;
Edwards, J. W. 2001b. Review of the forest habitat
relationships of the Indiana bat (Myotis sodalis).
Gen. Tech. Rep. NE-284. Newtown Square, PA: U.S.
Department of Agriculture, Forest Service,
Northeastern Research Station. 21 p.

Miller, G. W.; Wood, P. B.; Nichols. Two-age silviculture
—an innovative tool for enhancing species diversity
and vertical structure in Appalachian hardwoods.
In: Eskew, L. G., comp. Forest health through
silviculture. Proceedings of the 1995 national
silviculture workshop. 1995 May 8-11; Mescalero,
NM. Gen. Tech. Rep. RM-267. Fort Collins, CO:
U.S. Department of Agriculture, Forest Service, Rocky
Mountain Forest and Range Experiment Station: 175-
182.

Odom, R.H.; Ford, W. M.; Edwards, J. W.; Stihler, C.;
Menzel, J. M. 2001. Modeling Virginia northern
flying squirrel habitat in the central Appalachians.
Biological Conservation. 99: 245-252.

Owen, S. E; Menzel, M. A.; Ford, W. M.; Chapman, B.
R.; Miller, K. V.; Edwards, J. W.; Wood, P. B. 2001.
First summer record of a female Indiana bat, Myotis
sodalis, in West Virginia. Journal of the Elisha
Mitchell Scientific Society. 117: 132-134.

Racey, P. A. 1988. Reproductive assessment in bats. In:
Kunz, T. H., ed. Ecological and behavioral methods for
the study of bats. Washington, DC: Smithsonian
Institution Press: 31-45.

Rommé, R. C.; Tyrell, K.; Brack, V. 1995. Literature
summary and habitat suitability index model:
components of summer habitat for the Indiana bat,
Myotis sodalis. Fed. Aid Proj. E-1-7, Stud. No. 8.
Cincinnati, OH: 3D/Environmental. 38 p.

Sasse, D. B.; Pekins, P. J. 1996. Summer roosting
ecology of northern long-eared bats (Myotis
septentrionalis) in the White Mountain National
Forest. In: Barclay, R. M. R.; Brigham, R. M., eds.
Bats and forests symposium; 1995 October 19-21;
Victoria, BC. Victoria, BC: British Columbia Ministry
of Forests: 91-101.

Schuler, T. M; Gillespie, A. R. 2000. Temporal patterns
of woody species diversity in a central Appalachian
forest from 1856 to 1997. Journal of the Torrey
Botanical Society. 127: 149-161.

U.S. Department of Interior, Fish and Wildlife Service.
1999. Indiana bat (Myotis sodalis) revised recovery
plan. Fort Snelling, MN: U.S. Department of Interior,
Fish and Wildlife Service, Region 3. 53 p.

Vonhof, M. J.; Barclay, R. M. R. 1996. Roost-site
selection and roosting ecology of forest-dwelling
bats in southern British Columbia. Canadian
Journal of Zoology. 74: 1797-1805.

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

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