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BATS
IN FORESTS
This page intentionally left blank
BATS IN FORESTS
CONSERVATION AND MANAGEMENT
Edited by Michael J. Lacki, John P. Hayes, and Allen Kurta
Foreword by Merlin D. Tuttle

THE JOHNS HOPKINS UNIVERSITY PRESS

BALTIMORE
 This book was brought to publication
with the generous assistance of the
National Congress of American Indians,
Bat Conservation International,
and the Bat Diversity Network.

© 2007 The Johns Hopkins University Press

All rights reserved. Published 2007
Printed in the United States of America
on acid-free paper
1 2 3 4 5 6 7 8 9

The Johns Hopkins University Press

2715 North Charles Street
Baltimore, Maryland 21218-4363
www.press.jhu.edu

Library of Congress Cataloging-in-Publication Data

Bats in forests : conservation and management /
edited by Michael J. Lacki, John P. Hayes,
Allen Kurta.
p. cm.
Includes bibliographical references and index.
ISBN 0-8018-8499-3 (hardcover : alk. paper)
1. Bats—Ecology—North America. 2. Bats—
Effect of forest management on—
North America. 3. Bats—Conservation—
North America. I. Lacki, Michael J., 1956–
II. Hayes, John P. (John Parker)
III. Kurta, Allen, 1952–
QL737.C5B373 2007
639.97⬘94—dc22 2006013034

A catalog record for this book is available

from the British Library.

Photographs at chapter openers are by

Merlin D. Tuttle, Bat Conservation International
To Sonia, Ashley,
and Marty–
for always being there

To Joan, Heather,
and Forest

To my ‘Little Red Wife,’

for being
my best friend
This page intentionally left blank
 CONTENTS

Foreword, by Merlin D. Tuttle ix

Preface xi
Contributors xv

1 BATS IN FORESTS: WHAT WE KNOW

AND WHAT WE NEED TO LEARN
R. Mark Brigham 1
2 ECOLOGY AND BEHAVIOR OF BATS
ROOSTING IN TREE CAVITIES AND UNDER BARK
Robert M. R. Barclay and Allen Kurta 17
3 BEHAVIOR AND DAY-ROOSTING ECOLOGY
OF NORTH AMERICAN FOLIAGE-ROOSTING BATS
Timothy C. Carter and Jennifer M. Menzel 61
4 FORAGING ECOLOGY OF BATS IN FORESTS
Michael J. Lacki, Sybill K. Amelon, and Michael D. Baker 83
5 IMPORTANCE OF NIGHT ROOSTS TO THE ECOLOGY OF BATS
Patricia C. Ormsbee, James D. Kiser, and Stuart I. Perlmeter 129
6 MIGRATION AND USE OF AUTUMN, WINTER,
AND SPRING ROOSTS BY TREE BATS
Paul M. Cryan and Jacques P. Veilleux 153
7 SILVICULTURAL PRACTICES AND MANAGEMENT
OF HABITAT FOR BATS
James M. Guldin, William H. Emmingham, S. Andrew Carter,
and David A. Saugey 177
8 THE INFLUENCES OF FOREST MANAGEMENT
ON BATS IN NORTH AMERICA
John P. Hayes and Susan C. Loeb 207
9 ECOLOGICAL CONSIDERATIONS FOR LANDSCAPE-LEVEL
MANAGEMENT OF BATS
Joseph E. Duchamp, Edward B. Arnett, Michael A. Larson,
and Robert K. Swihart 237
10 ASSESSING POPULATION STATUS OF BATS IN FORESTS:
CHALLENGES AND OPPORTUNITIES
Theodore J. Weller 263
11 PLANNING FOR BATS ON FOREST INDUSTRY LANDS IN
NORTH AMERICA
T. Bently Wigley, Darren A. Miller, and Greg K. Yarrow 293

Author Index 319

Species Index 325
Subject Index 327
This page intentionally left blank
FOREWORD

Today, many bat populations are only small remnants of former num-
bers. Prior to the arrival of European settlers in the United States, for ex-
ample, millions of now endangered Indiana and gray myotis lived in sin-
gle caves. Even species that are still considered common have, in fact,
declined markedly. For example, in the 1870s great migratory flocks of
eastern red bats were reported to pass over for days at a time during fall
migration, a sight now vanished from American skies.
Despite their decline and an increase in conservation concern, bats
still rank among the world’s most neglected and misunderstood mam-
mals. Few members of the public understand that the world’s more than
1,100 species play essential ecological roles in all but the most extreme
desert and polar regions. Fewer still know that most of America’s 51
species rely on forests to varying degrees, either for food or shelter. In
turn, forests nearly everywhere benefit from bats, probably much more
than is realized. The most abundant bats inhabiting forests consume vast
quantities of insects nightly, including costly “pests” such as moths and
beetles. In short, the available evidence suggests that bats are essential to
ecological balance and forest health, fulfilling the same roles by night as
birds do by day.
A variety of factors related to forest conditions have contributed to bat
decline, including disturbance of hibernation and nursery caves in
forests, loss of snags and major cavities in ancient trees, careless use of
pesticides in forest settings, and early neglect of forest management. For-
est bats often require loose, exfoliating bark and other cavities in extra
tall trees that are dying or newly dead, and these are now scarce in many
places. Bats also require alternative roosts in close proximity, in part be-
cause exfoliating bark and old snags are ephemeral in nature, and in part
because they must find varied temperatures and avoid predators.
Management practices that ensure age diversity of trees, retain snags
and damaged trees, and aid plant diversity can greatly benefit bat popu-
lations. When forests become monocultures, bats and other insectivo-
rous animals encounter unfavorable conditions such as prey cycles that
produce fewer species of insects and longer gaps between hatch cycles.
The eventual explosions of these insects then results in their designation
as major forest pests. Both forests and bats are more secure when struc-
tural, floral, and faunal diversity are maintained. For these reasons, the
 x growing collaboration between bat biologists and forestry agencies and
Foreword industries is a wise investment.
In this book, America’s leading experts on bats collaborated to pro-
duce an extraordinarily comprehensive review of current knowledge that
is the best available anywhere. Both the volume editors and chapter au-
thors are unsurpassed in their respective areas. Twenty-seven authors
have contributed eleven chapters of up-to-date research and review that
is a must read for anyone interested in either bats or forest management.

Merlin D. Tuttle
PREFACE

At least 45 species of bats occur in the United States and Canada, and 27
of these use trees for roosting at least some of the time (see table P.1), and
presumably forage in or near forested areas as well. All forest-dwelling
bats in North America are insectivorous, and all except one species, the
pallid bat (Antrozous pallidus; Antrozoidae), belong to the family Ves-
pertilionidae. Many species of bats that inhabit forests are species of con-
cern, threatened, or endangered, because of low or unknown population
size, apparent population declines, high vulnerability to disturbance, or
limited geographic distribution. Although some species, such as the lit-
tle brown myotis (Myotis lucifugus) and big brown bat (Eptesicus fuscus),
have been studied for more than 50 years, there is surprisingly little in-
formation on how these or other bats use forested areas. In fact, some
species of bats are so poorly studied that current approaches to conser-
vation and management of these species within forests is reduced to an
educated guess. The recent spate of lawsuits to halt logging on public
lands in the eastern United States, due largely to the endangered Indiana
myotis (Myotis sodalis), emphasizes the need for better information on
bats in forests.
Data on forest-dwelling bats are mostly scattered throughout scien-
tific journals and the immense gray literature of governmental agencies
and private industry. To date, only two consolidated volumes of infor-
mation on forest-dwelling bats in North America have been published
(R. M. R. Barclay and R. M. Brigham, 1996, Bats and Forests Symposium;
A. Kurta and J. Kennedy, 2002, The Indiana Bat: Biology and Management
of an Endangered Species). In both books, however, individual chapters
are limited in scope and, in general, present new data on a specific topic,
rather than synthetic reviews of existing data or development of strate-
gies for management of bats and their habitats.
The First Bats and Forests Symposium held in 1995 in Victoria, British
Columbia, was an initial attempt to bring together individuals engaged
in the study or management of forest-dwelling bats in the United States
and Canada. Since then, advances in technologies and approaches to col-
lecting data, including reduction in mass of radiotransmitters and im-
provement of acoustic-monitoring systems, have led to new insights into
the biology of many species of forest-dwelling bats. Moreover, the num-
ber of biologists, consultants, and land managers involved with bats in
 xii forests has increased in recent decades, resulting in a large volume of both
Preface published and unpublished data and a growing need for synthesis of ex-
isting information. For example, of the papers presented at the North
American Symposium on Bat Research from 1991 to 1993, 24 (or 9.3%
of the total presentations) addressed topics on North American forest-
dwelling bats, whereas from 2001 to 2003, 71 papers (19.1%) were asso-
ciated with bats in forests. This increase in attention to forest-dwelling
bats is evident in other scientific outlets as well. From 1986 to 1991, the
Journal of Wildlife Management published only a single article addressing
bats in forests, but from 1999 to 2004 the number of articles on bats in
North American forests had increased to 12. Additionally, articles ad-
dressing management of forest-dwelling bats are now appearing in
forestry journals, such as Forest Science, Forest Ecology and Management,
and Northern Journal of Applied Forestry; outlets in which data on bats
and their habitats had previously not been published.
To help synthesize new information on bats in forests and provide a
foundation for future directions in the field, the Second Bats and Forests
Symposium and Workshop was held in Hot Springs, Arkansas, in 2004.

Table P.1. Regional distribution of occurrence of North American

tree-roosting bats

Distribution in
Species Common name North America
Antrozous pallidus pallid bat NW, SW
Corynorhinus rafinesquii Rafinesque’s big-eared bat SE
Corynorhinus townsendii Townsend’s big-eared bat NW, SE, SW
Eptesicus fuscus big brown bat All
Euderma maculatum spotted bat NW, SW
Idionycteris phyllotis Allen’s big-eared bat SW
Lasionycteris noctivagans silver-haired bat All
Lasiurus blossevillii western red bat NW, SW
Lasiurus borealis eastern red bat All
Lasiurus cinereus hoary bat All
Lasiurus ega southern yellow bat SW
Lasiurus intermedius northern yellow bat SE
Lasiurus seminolus Seminole bat SE
Lasiurus xanthinus western yellow bat SW
Myotis auriculus southwestern myotis SW
Myotis austroriparius southeastern myotis SE
Myotis californicus California myotis NW, SW
Myotis evotis long-eared myotis NW, SW
Myotis keenii Keen’s myotis NW
Myotis lucifugus little brown myotis All
Myotis septentrionalis northern myotis NE, NW, SE
Myotis sodalis Indiana myotis NE, SE
Myotis thysanodes fringed myotis NW, SW
Myotis volans long-legged myotis NW, SW
Myotis yumanensis Yuma myotis NW, SW
Nycticeius humeralis evening bat NE, SE, SW
Pipistrellus subflavus eastern pipistrelle NE, SE, SW
This meeting was built around a series of invited presentations that re- xiii
viewed our scientific understanding of key issues related to the ecology Preface
and management of bats in North America forests. The presentations
were followed by workshops and field tours that brought together biolo-
gists and forest managers to help focus future research directions and im-
prove strategies for management of bat habitat. The present book, Bats
in Forests: Conservation and Management, grew from the seeds planted
at that meeting and was written to meet the need for a synthesis of in-
formation on forest-dwelling bats in North America.
The book is structured around 11 chapters, beginning with an intro-
duction to the topic of bats in North American forests that highlights key
areas of progress and recommended avenues for future study of forest-
dwelling bats. This chapter is followed by a series of papers that discuss
use of roost trees, including cavity, bark, and foliage roosts, foraging habi-
tat, and night roosts of bats. The final section of the book includes chap-
ters that address silviculture and forest management, including manage-
ment for bats on private industry lands, monitoring and censusing of bat
populations, and response of bats to forest management at the level of
the stand and landscape.
Authors of chapters were chosen for their expertise in particular ar-
eas, and, to ensure breadth and depth of coverage, most chapters have
multiple authors representing different regions of the continent and/or
areas of expertise. This range of backgrounds and knowledge provides
the reader with differing perspectives regarding bats in forests. The book
is intended to serve as a resource for students, academic biologists, gov-
ernmental employees, environmental organizations, biological consul-
tants, legal experts, and private land managers. The emphasis of this book
is on bats in North American forests, so the literature cited and data sets
evaluated primarily represent studies on North American species; but
this volume should also serve as a comparative reference for anyone con-
cerned with forest-dwelling bats in other temperate regions of the globe
because many of the problems associated with management of bat habi-
tat are the same.
Each manuscript was peer-reviewed to facilitate a more comprehen-
sive coverage of material and to set a standard of quality in presentation
of each chapter. We thank the following individuals for reviewing vari-
ous chapters: Ed Arnett, Mike Baker, Burr Betts, Mark Brigham, Carol
Chambers, John Cox, Paul Cryan, Mark Ford, Matina Kalcounis-Rüppell,
Steve Langenstein, Jeff Larkin, Susan Loeb, Alice Chung-MacCoubrey,
Chester Martin, Darren Miller, Susan Murray, Ralph Nyland, Tom
O’Shea, Dan Taylor, Ted Weller, and Craig Willis. Financial assistance
with both the Symposium and Workshop and with the preparation of
this book was provided by Bat Conservation International, Inc., Weyer-
haeuser Company, National Council for Air and Stream Improvement,
Inc., Southeastern Bat Diversity Network, and the College of Agriculture
at the University of Kentucky. We thank Dan Taylor and Merlin Tuttle for
their assistance with development of the Symposium and Workshop and
for support in completing this project. The editors also wish to recognize
 xiv our home institutions, the University of Kentucky, University of Florida,
Preface and Eastern Michigan University for infrastructural and financial sup-
port of our research programs over the past two decades, and to all the
sponsors, federal, state, and private, that have contributed technical and
logistical support, and financial resources to assist us in the study of bats
in forests.
CONTRIBUTORS

S K . A, Research Wildlife Biologist, North Central Research

Station, University of Missouri, Columbia, Missouri 65211
E  B. A , Conservation Scientist, Bat Conservation Interna-
tional, Inc., Austin, Texas 78746
M D. B, Post-doctoral Scholar, Department of Forestry, Uni-
versity of Kentucky, Lexington, Kentucky 40546
R M. R. B , Professor, Department of Biological Sciences,
University of Calgary, Calgary, Alberta, Canada T2N 1N4
R. M B, Professor, Department of Biology, University of
Regina, Regina, Saskatchewan, Canada S4S 0A2
S. A C, Wildlife Biologist, Southern Research Station, USDA
Forest Service, Nacogdoches, Texas 75965
T  C. C , Associate Scientist, Department of Zoology, South-
ern Illinois University, Carbondale, Illinois 62901
P  M. C , Wildlife Biologist, USGS, Fort Collins Science Center, Fort
Collins, Colorado 80526
J E . D, Doctoral Candidate, Department of Forestry and
Natural Resources, Purdue University, West Lafayette, Indiana 47907
W H. E, Emeritus Professor, Department of Forest Sci-
ence, Oregon State University, Corvallis, Oregon 97331
J M. G, Supervisory Ecologist, Southern Research Station,
USDA Forest Service, P.O. Box 1270, Hot Springs, Arkansas 71902
J P. H , Professor & Chair, Department of Wildlife Ecology and
Conservation, University of Florida, Gainesville, Florida 32611
J D. K, Wildlife Ecologist, Daniel Boone National Forest, Whitley
City, Kentucky 42653
A K, Professor, Department of Biology, Eastern Michigan Univer-
sity, Ypsilanti, Michigan 48197
M J. L , Professor, Department of Forestry, University of Ken-
tucky, Lexington, Kentucky 40546
M A. L, Wildlife Research Scientist, Forest Wildlife Research
Group, Minnesota Department of Natural Resources, Grand Rapids, Min-
nesota 55744
S C. L, Wildlife Ecologist, Southern Research Station, Department
of Forest Resources, Clemson University, Clemson, South Carolina 29634
J M. M, Research Wildlife Biologist, Northeastern Research
Station, USDA Forest Service, Parsons, West Virginia 26287
 xvi D A. M, Southern Wildlife Program Manager, Weyerhaeuser
Contributors Company, Columbus, Mississippi 39704
P C. O, Wildlife Ecologist, Willamette National Forest,
Eugene, Oregon 97440
St I. P, WELL Project Coordinator, Springfield Public
Schools, Springfield, Oregon 97477
D A. S, Wildlife Biologist, Ouachita National Forest, Jessieville,
Arkansas 71949
R K. S, Professor & Interim Head, Department of Forestry
and Natural Resources, Purdue University, West Lafayette, Indiana 47907
J P. V, Assistant Professor, Department of Biology, Franklin
Pierce College, Rindge, New Hampshire 03461
T  J. W, Research Wildlife Ecologist, Pacific Southern
Research Station, USDA Forest Service, Arcata, California 95521
T. B W, Manager of the Sustainable Forestry and Eastern
Wildlife Program, National Council for Air and Stream Improvement, Inc.,
Clemson, South Carolina 29634
G K. Y, Professor, Department of Forestry and Natural
Resources, Clemson University, Clemson, South Carolina 29634
BATS
IN FORESTS
BATS IN FORESTS: WHAT WE KNOW
AND WHAT WE NEED TO LEARN
R. Mark Brigham
1

Ten years ago, reports about the ecology, management, and behavior of
bats in North America were, with few exceptions, centered on aggrega-
tions of animals in caves and human-made structures. Over the past
decade, however, the focus has changed toward trying to understand how
many of these same species interact with forested environments and how
we need to manage forests with a view to conserving the bats inhabiting
them. Hayes (2003) recently pointed out that the degree of association
between bats and forests varies considerably and that the degree of asso-
ciation represents a continuum rather than a set of discrete categories. Of
the 45 species of bats recognized in North America (O’Shea and Bogan
2003), 25 species use forests for roosting or foraging at least some of the
time. Most “forest-dwelling bats” are small (⬍30 g), and virtually all are
aerial insectivores, although some glean invertebrates from the ground
or the surface of vegetation (Faure et al. 1993; Faure and Barclay 1992;
Lacki et al., chap. 4 in this volume). A significant portion of the bat com-
munity in North America is associated with wooded areas, whether for
foraging, roosting, or both. Thus, it is important to consider the nature
of the interaction between these animals and forested habitats.
This chapter provides a brief synthetic perspective of the general fea-
tures of what biologists know about forest-dwelling bats in North Amer-
ica, most notably information published in the past decade, and it offers
some opinion about the priorities for future studies. My review is by no
means exhaustive, and thorough assessments of various aspects of the
lives of forest-dwelling bats can be found in Hayes (2003), Miller et al.
(2003), and subsequent chapters in this volume. This chapter is North
American in outlook, but I encourage readers to examine recent studies
of bats emanating from New Zealand (O’Donnell and Sedgeley 1999;
Sedgeley and O’Donnell 1999), Australia (Law and Chidel 2002; Lums-
den et al. 2002; Pavey and Burwell 2004; Turbill et al. 2003), and Europe
(Kerth and König 1999; Russo et al. 2002; Russo and Jones 2003). Al-
though there may be geographic differences, these studies provide a valu-
able ecological corollary to North American research, and it would be a
grave mistake to adopt a continentally insular perspective.

ROOSTS
Roosts are critically important for bats because they provide shelter from
the elements and a haven from potential predators, as well as a location
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