United States

Department of

Agriculture

Forest Service

Northeastern

Research Station

General Technical

Report NE-292

Roost Tree Selection by

Maternal Colonies of Northern

Long-eared Myotis in an

Intensively Managed Forest

Sheldon F. Owen

Michael A. Menzel

W. Mark Ford

John W. Edwards

Brian R. Chapman

Karl V. Miller

Petra B. Wood
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Abstract

We attached radio transmitters to 20 pregnant or lactating northern long-eared

myotis (Myotis septentrionalis) and located 43 maternity colony roost trees in

intensively managed hardwood forests in the Allegheny Mountains of east-central

West Virginia. Maternity colonies ranged in size from 7 to 88 individuals. We

compared 23 characteristics of the 43 roosts to 43 randomly selected trees with

cavities or exfoliating bark. Roost trees selected by maternity colonies differed from

randomly located roost trees in that they were taller, smaller in diameter, surrounded

by more live overstory trees and snags, and surrounded by a higher basal area of

other snags. Black locust (Robinia pseudoacacia) and black cherry (Prunus

serotina) were used as day roosts more frequently than expected based on their

abundance across this landscape. Intensively managed hardwood forests in the

central Appalachians apparently provide adequate roosting habitat for northern longeared myotis.

The Authors

SHELDON F. OWEN and MICHAEL A. MENZEL are graduate research assistants in

the Division of Forestry, West Virginia University in Morgantown. W. MARK FORD is

a research wildlife biologist with the USDA Forest Service Northeastern Research

Station in Parsons, West Virginia. JOHN W. EDWARDS is an assistant professor in

the Division of Forestry, West Virginia University. BRIAN R. CHAPMAN is the dean

of College of Arts and Sciences at Sam Houston State University in Huntsville,

Texas. KARL V. MILLER is a professor in the Warnell School of Forest Resources,

University of Georgia in Athens. PETRA B. WOOD is the assistant unit leader for the

USGS Biological Resource Division’s West Virginia Cooperative Fish and Wildlife

Research Unit in Morgantown.

Manuscript received for publication 19 October 2001

Cover

The Shawnee National Forest is a nesting site for the northern long-eared bat

(photo by Timothy C. Carter, Southern Illinois University).
1

Introduction

Many North American bat species reside in forests and

depend on living trees or snags for roosting sites

(Barbour and Davis 1969, Humphrey et al. 1977, Kunz

1982, Vonhof and Barclay 1996). Roosts provide

thermally stable environments and protection from the

elements and predators (Kunz 1982). Initial research in

roosting ecology focused on easily accessible manmade

structures or caves (Kunz 1982, Betts 1996). Most early

records of tree roost sites were anecdotal descriptions of

single-tree colonies of bats (Humphrey et al. 1977, Kunz

1982, Barclay and Cash 1985). However, since the

advent of miniaturized radio transmitters suitable for

bats, there has been a renewed interest in roost ecology

research (Kunz 1982, Cambell et al. 1996, Sasse and

Perkins 1996, Vonhof and Barclay 1996, Foster and

Kurta 1999).

In the western United States, old-growth forests are

important bat roosting habitats (Thomas 1988,

Ruggiero et al. 1991, Campbell et al. 1996, Zielinski and

Gellman 1996). However, in the central Appalachians,

extensive timber harvesting in the early 1900s and the

more recent increase in forest harvesting over the last

decade have resulted in many high-graded stands with

80- to 100-year-old unmerchantable residual trees and

frequent canopy caps or even-aged clearcut stands 20

years old or younger (McGarigal and Fraser 1984,

Rosenberg et al. 1988, Weakland 2000). Characteristics

of snags and cavity trees in these intensively utilized

forests differ from those in less intensively managed

(locally, stands not entered since the early 1900s) or

old-growth stands (McComb and Muller 1983,

Rosenberg et al. 1988). Regionally, remaining oldgrowth stands are too small and disjunct to be

biologically meaningful for bats.

Roost habitat destruction and alteration is thought to be

one of the most detrimental factors influencing bats in

eastern North America (Kunz 1982, Menzel et al. 2001).

An understanding of roost requirements of common

species, such as the northern long-eared myotis, is vital

to bat conservation in the central Appalachians and

elsewhere. Our study objectives were to determine if and

where northern long-eared myotis maternity colonies

roosted in intensively managed central Appalachian

forest stands and to compare roost trees (and the

surrounding habitat) with potential roost trees to

identify important roost habitat characteristics.

Study Area

Our study was centered on the Westvaco Wildlife and

Ecosystem Research Forest (WERF) and surrounding

Westvaco Forest Resources properties. The WERF is a

3,360-ha area located in the unglaciated Allegheny

Mountain and Plateau physiographic province of West

Virginia (Fenneman 1938) in southwestern Randolph

County (38° 42' N lat., 80° 3' W long.). Established by

Westvaco Corp. in 1994, the area is reserved for the

study of industrial forestry impacts on ecosystems and

ecological processes in an Appalachian setting. Oldest

forests on the WERF are second-growth stands that were

established by natural regeneration following the

extensive railroad logging that occurred in this portion

of West Virginia from 1900 to 1930 (Clarkston 1993).

Elevations range from 740 to 1200 m. Topography

consists of steep side slopes with broad, plateau-like

ridgetops and narrow valleys with small, high-gradient

streams. The climate is cool and moist with average

annual precipitation exceeding 155 cm (Strausbaugh

and Core 1977). Soils on the area are classified as acidic

and well-drained Inceptisols (Stephenson 1993).

Overstory forest cover is primarily an Allegheny

hardwood-northern hardwood type dominated by beech

(Fagus grandifolia), yellow birch (Betula alleghaniensis),

sugar maple (Acer saccharum), red maple (A. rubrum),

black cherry (Prunus serotina), Fraser’s magnolia

(Magnolia fraseri) and red spruce (Picea rubens). Species

from the cove hardwood and mixed mesophytic

associations, including yellow-poplar (Liriodendron

tulipifera), basswood (Tilia americana), sweet birch (B.

lenta), and northern red oak (Quercus rubra) occur at

lower elevations. Overtopped and suppressed or dead

black locust (Robinia pseudoacacia) trees and snags are

common in many forest stands. Riparian areas are

characterized by eastern hemlock (Tsuga canadensis)-

rhododendron (Rhododendron maximum) communities.

In much of the upland area, rhododendron and striped

maple (A. pensylvanicum) form dense understory

thickets. An artifact of past forest management practices

and white-tailed deer (Odocoileus virginianus) herbivory,

a dense cover of hay-scented fern (Dennstaedtia

punctilobula) occurs where the shrub layer is absent and

the overstory canopy is not continuous (Ford and

Rodrigue 2001). Recent (5 years old or younger)

selection harvests and clearcuts are common on the area.

Where regeneration is successful following harvest,

stands are dominated by copious amounts of red maple,

yellow birch, black cherry, fire cherry (Prunus

allegheniensis), yellow poplar, American beech and black

locust stems.

Materials and Methods

We captured northern long-eared myotis from June to

August 1998 and from May to August 1999 using 12 x

2.4 m mist nets and high-net systems consisting of three

nets stacked vertically (Gardner et al. 1989). Nets were

positioned over streams, ephemeral pools, road ruts,

and water-filled borrow pits. We defined a mist-net

night as a single mist net open for at least 1 hour. Data

collected from each captured bat included species, age,

sex, mass, forearm length, and reproductive condition.

We determined female reproductive condition by

palpating the abdomen and inspecting the mammae

and determined age-class by examining the degree of
2

epiphyseal-diaphyseal fusion (Anthony 1988, Racey

1988).

We attached LB-2 radio transmitters (0.46 and 0.55 g;

Holohil Systems Ltd., Woodlawn, ON) to 20 pregnant

or lactating northern long-eared myotis. Transmitters

were attached to the hair between the scapulae using

Skin Bond® surgical adhesive (Pfizer Hospital Products

Group, Inc., Largo, FL). Transmitters weighed less than 5

percent of any individual bat’s body weight (Aldridge

and Brigham 1988) and each had a battery life of

approximately 21 days.

We located roosting northern long-eared myotis during

the day using standard radiotelemetry techniques. We

used R2000 ATS receivers (Advanced Telemetry Systems,

Inc., Isanti, MN) and folding, three-element Yagi

antennas to return to trees that evening to determine the

exact location of roosts and relative sizes of the maternal

colonies. All bats were tracked until they dropped the

transmitter or the battery failed ( × = 6 days; SE =2.5; n

= 20).

To identify habitat features that might influence roost

selection, we measured 23 characteristics of the roost

tree and surrounding habitat. We compared these

measurements to data from apparently suitable but

unused roost trees that contained either a cavity or large

plates of exfoliating bark (Betts 1997). We located

random suitable trees by generating a set of Universal

Transverse Mercator coordinates within 0.5 km of an

actual roost tree. From that location, we walked

concentric circles until we found a suitable tree with a

cavity or exfoliating bark. We subjectively assessed the

level of structural complexity within 1 m2 directly in

front of the portion of the tree considered the roost-area

entrance. A complexity value of 1 indicated that there

was no vegetation in front of the roost; a value of 5

indicated that the roost entrance was obscured by dense

vegetation. We also measured height of the roost

entrance, total tree height, and diameter at breast height

(d.b.h.).

We measured 18 variables within 0.04-ha plots centered

on the roost tree (Table 1). We recorded the distance

from the roost tree to the nearest overstory tree and to

the nearest taller overstory tree. Canopy density was

measured using a spherical densiometer (Lemmon

1956) at 2 m from the bole of the roost tree on the same

side as the roost. Overstory trees were defined as greater

than 9.5 cm d.b.h, whereas understory trees were woody

stems between 1.0 and 9.5 cm d.b.h. The percentage of

overstory trees composed of conifers or snags was based

on basal area. Shannon’s measure of diversity, Pielou’s

measure of evenness, and simple species richness were

used for both overstory and understory trees in these

analyses (Pielou 1966). Understory density was

recorded as the number of woody stems less than 9.5

cm d.b.h. within the 0.04-ha plot.

Characteristics of roost trees and random trees were

compared using two sample t-tests testing a two-tailed

alternative hypothesis of unequal means (SAS Institute

1990). Significance was set at alpha = 0.05. We recorded

all trees by species along 12 randomly located transects

( × = 153.0 m; SE = 31.9) to further evaluate tree species

selection by comparing roost trees to the available tree

species with possible roost sites (snags or live trees with

cavities or exfoliating bark). We then compared the

proportion of tree species used as roost trees to the

proportion of available roost tree species using Chisquare analysis (SAS Institute 1990).

Results

We captured 168 bats of 8 species in 221 mist-net

nights. These included 23 big brown bats (Eptesicus

fuscus), 3 silver-haired bats (Lasionycteris noctivagans), 25

eastern red bats (Lasiurus borealis), 12 hoary bats

(Lasiurus cinereus), 27 little brown bats (Myotis lucifugus),

61 northern long-eared myotis (M. septentrionalis), one

Indiana bat (M. sodalis), and 16 eastern pipistrelles

(Pipistrellus subflavus).

We located 43 roost trees used by northern long-eared

myotis. Females changed roost location an average of

every three days ( × = 3; SE = 1.5). Many bats used the

same roost tree simultaneously or over successive days.

Based on our emergence counts, size of maternity

colonies ranged from 7 to 88 individuals. On 22 June

1999, we counted 88 northern long-eared myotis exiting

a cavity in a live black cherry. We found 15 roosts under

exfoliating bark, whereas 28 were found in natural

cavities.

The mean height of actual roosts was higher (P = 0.003)

than that of the potential roost trees (Table 1). The

complexity surrounding the roost entrance did not differ

(P = 0.4081) between the actual roost and potential

roosts. Although d.b.h. of roost trees was smaller (P =

0.0249) than that of potential roost trees, roost tree

d.b.h. did not differ from that of other trees within the

plot. Only 3 of the 18 site characteristics differed

between roost and random sites (Table 1). The basal

area of snags and percentage of snags in the overstory

were higher surrounding roost trees than random trees

(P = 0.0425; P = 0.0376). The number of stems in the

overstory also was higher around roost trees than the

potential roost trees (P = 0.0403).

Roosts were located in 11 tree species (Table 2). We

found 13 (30 percent) roosts in black locust and nine

(21 percent) in black cherry. Black locust comprised 9

percent of the available trees and black cherry 11 percent

of the available trees. Black locust and black cherry were

used more often than expected relative to the proportion

available (χ2 = 35.59, 9 df, P < 0.001).
3

Table 1.—Comparison of means (+ SE) of 23 characteristics between 43 roost trees used by maternity colonies of

Myotis septentrionalis and 43 potential roost trees using two sample t-tests.

Roost Potential

Mean SE Mean SE t P

Cavity characteristics

Roost height (m) 10.8 0.6 7.7 0.6 -3.78 0.0003

Complexity 1.8 0.2 1.6 0.1 -0.83 0.4081

Tree characteristics

Height of roost tree (m) 17.8 0.7 15.9 1.0 -1.63 0.1074

Diameter of roost tree (cm) 27.2 1.0 31.1 1.4 2.28 0.0249

Roost d.b.h. vs mean d.b.h. of plot 26.3 1.3 24.3 0.7 -1.46 0.1500

Site characteristics

Nearest overstory tree (m) 1.6 0.2 2.1 0.4 1.13 0.2632

Nearest taller overstory tree (m) 2.8 0.4 3.6 0.6 1.19 0.2371

Difference in roost and canopy tree height -1.6 1.8 -5.6 1.9 -1.52 0.1331

Percent canopy density 90.9 2.2 86.1 3.6 -1.14 0.2589

Degree of slope 18.0 1.7 19.0 1.2 0.57 0.5676

Number of overstory trees 24.8 1.3 20.7 1.5 -2.08 0.0403

Total basal area of plot 13.8 0.6 13.0 0.8 -0.76 0.4471

Basal area of snags 0.9 0.2 0.5 0.1 -2.07 0.0425

Percent snags in overstory 6.9 1.4 3.7 0.6 -2.13 0.0376

Conifer basal area 1.4 0.3 1.1 0.3 -0.54 0.5883

Percent conifers in overstory 9.9 2.3 8.6 2.1 -0.43 0.6695

Overstory richness 6.7 0.31 6.1 0.35 -1.29 0.1999

Overstory diversity 1.5 0.06 1.5 0.08 -0.25 0.800

Overstory evenness 0.8 0.02 0.8 0.04 -0.05 0.9602

Density of understory 9.7 1.5 8.5 1.0 -0.67 0.5067

Understory richness 2.4 0.20 2.4 0.22 0.00 1.000

Understory diversity 0.6 0.07 0.6 0.08 0.17 0.8626

Understory evenness 0.6 0.1 0.6 0.1 0.08 0.9397

Table 2.—Tree species chosen as roosts for maternity colonies by Myotis septentrionalis

compared to available potential roost tree species.

Species Maternal roosts Percent Potential roosts Percent

Robinia pseudoacacia 13 30.2 27 9.0

Prunus serotina 9 20.9 32 10.7

Acer spp. 7 16.3 37 12.3

Quercus spp. 5 11.6 22 7.3

Magnolia spp. 4 9.3 25 8.3

Other 3 7.0 54 18

Fagus grandifolia 1 2.3 28 9.3

Betula spp. 1 2.3 50 16.7

Lirodendron tulipifera 0 0.0 13 4.3

Fraxinus americanus 0 0.0 12 4.0
4

Discussion

Similar to our results, Sasse and Perkins (1996) reported

36 northern long-eared myotis leaving a tree cavity in

northern New Hampshire, whereas Foster and Kurta

(1999) counted 60 individuals from a roost in

Michigan. We also counted 77 and 64 bats from two

other roosts within 1 km of the 88-exit count. We

believe the large colony sizes we observed indicate that

intensively managed forests in the central Appalachian

do provide suitable habitat for northern long-eared

myotis.

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). Presumably, these

larger trees have increased solar exposure that aids in

juvenile growth and development (Menzel et al. 2001).

The aforementioned studies sharply contrast with this

research indicating that northern long-eared myotis

tended to select smaller trees in more cluttered stands.

Moreover, our data challenges the assumption that in

cool-temperate environments, tree-roosting bats will

select roost to optimize solar exposure. We believe our

most important discovery is that northern long-eared

myotis roosted in localized areas with abundant snags.

Whether northern long-eared myotis choose these sites

and roosts from ease of initial discovery or the ability to

have multiple alternate roosts in a more compact area is

unknown.

We caution that lower roost heights for the potenial

roost trees may be biased due to increased observer

visibility, but northern long-eared myotis are small, very

agile bats with low wing loading and aspect ratio with

gleaning food habits well adapted to feed in or around

vegetation (Barbour and Davis 1969, Faure et al. 1993,

Findley 1993). Our roost data supports the hypothesis

of Foster and Kurta (1999) that the increased agility

allows northern long-eared myotis to easily exploit

roosts in cluttered environments.

Across their distribution, northern long-eared myotis

roost in many tree species. Mumford and Cope (1964)

and Clark et al. (1987) found northern long-eared

myotis roosting in elm (Ulmus spp.) in Indiana and

Iowa. Sasse and Perkins (1996) observed roosts in

American beech and sugar maple in New Hampshire.

Working in Michigan, Foster and Kurta (1999) were the

first to report silver maple (Acer saccharinum) and green

ash (Fraxinus pennsylvanica) used as roosts. Our research

is the first to identify the use of black locust and black

cherry as roosts for this bat species. Both of these tree

species were used in greater proportion than the relative

proportion available in the surrounding forest stands.

Black locust is an intolerant pioneer species in the

central Appalachians with quick regenerative response

after a ground-clearing disturbance such as timber

harvest. The species initially exhibits rapid growth, but is

soon overtopped and suppressed by other more longlived tree species. These overtopped and suppressed

black locust usually die, leaving a rot-resistant snag that

can remain standing for years. Black cherry also is an

intolerant species that responds to complete overstory

disturbance and removal by exhibiting rapid growth

(Smith 1994, Harlow et al. 1996). Regionally, black

cherry regenerates in such large numbers that high

numbers of snags are inevitably available throughout

most times during a forest stand’s development (70- to

120-year rotation for sawtimber; 40- to 70-year rotation

for fiber production). Black cherry also is resistant to rot

and can produce large, quality roost snags. Our results

support the hypothesis of Thomas (1988) that snag

longevity is an important factor in bat roost selection.

Based on our findings, we believe roost availability for

northern long-eared myotis is not a biologically limiting

factor in intensively managed forests in the central

Appalachians.

Acknowledgements

Financial support for this project was provided by

Westvaco Corporation’s Ecosystem Research Forest

Program, USDA Forest Service Monongahela National

Forest, University of Georgia Warnell School of Forest

Resources, and West Virginia University Division of

Forestry. Field assistance was provided by J. Johnson, D.

Leput, B. Smith, and J. Menzel. An earlier draft of this

paper was reviewed by J. Carroll and S. Cross.

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Printed on Recycled Paper

Owen, Sheldon F.; Menzel, Michael A.; Ford, W. Mark; Edwards, John W.;

Chapman, Brian R.; Miller, Karl V.; Wood, Petra B. 2002. Roost tree selection

by maternal colonies of northern long-eared myotis in an intensively

managed forest. Gen. Tech. Rep. NE-292. Newtown Square, PA: U.S.

Department of Agriculture, Forest service, Northeastern Research Station. 6 p.

Presents data on the tree-roosting ecology of female northern long-eared myotis

(Myotis septentrionalis) in managed hardwood forests in the Allegheny Mountains of

West Virginia.

Keywords: black locust, cavities, Myotis septentrionalis, northern long-eared

myotis, Robinia pseudoacacia, roost snag
Headquarters of the Northeastern Research Station is in Newtown Square,

Pennsylvania. Field laboratories are maintained at:

Amherst, Massachusetts, in cooperation with the University of Massachusetts

Burlington, Vermont, in cooperation with the University of Vermont

Delaware, Ohio

Durham, New Hampshire, in cooperation with the University of New Hampshire

Hamden, Connecticut, in cooperation with Yale University

Morgantown, West Virginia, in cooperation with West Virginia University

Parsons, West Virginia

Princeton, West Virginia

Syracuse, New York, in cooperation with the State University of New York,

College of Environmental Sciences and Forestry at Syracuse University

Warren, Pennsylvania

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