Ecology

of Bats

This limited facsimile editian has been issued

for the purpose of keeping this title avaiJable
to the scientific community.

Ecology
of Bats
Edited by

Thomas H. Kunz
Boston University
Boston, Massachusetts

Plenum Press New York and London

Library of Congress Cataloging in Publication Data

Main entry under title:
Ecology of bats.
Indudes bibliographical references and index.
1. Bats-Ecology. 2. MammaIs-Ecology. I. Kunz, Thomas H.
QL737.C5E33 1982
599.4'045
ISBN-13: 978-1-4613-3423-1
001: 10.1007/978-1-4613-3421-7

82-10157

e-ISBN-13: 978-1-4613-3421-7

AACR2

This limited facsimile edition has been issued

for the purpose of keeping this title available
to the scientific community.

10987654

1982 Plenum Publishing Corporation

Softcover reprint of the hardcover 1st ed ition 1982

233 Spring Street, New York, N.Y. 10013

All rights reserved
No part of this book may be reproduced, stored in a retrieva! system, or transmitted,
in any form or by any means, electronic, mechanieal, photocopying, micromming,
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To Margaret, Pamela, and David

Contributors
Hans G. Erkert Institut fr Biologie III der Universitt Tbingen. D-7400
Tbingen I, Federal Republic of Germany
M. Brock Fenton Department of Biology. Carleton University, Ottawa KIS
5B6. Canada
James S. Findley Department of Biology, University of New Mexico. Albuquerque. New Mexico 87131
Theodore H. Fleming Department of Biology. University of Miami, Coral
Gables. Florida 33124
E. Raymond Heithaus Department of Biology, Kenyon College. Gambier.
Ohio 43022
Thomas H. Kunz Department of Biology, Boston University, Boston, Massachusetts 02215
Adrian G. Marshall Department of Zoology, University of Aberdeen. Aberdeen AB9 2TN. Scotiand
Brian K. McNab Department of Zoology. University of Florida, Gainesville,
Florida 32611
P. A. Racey Department of Zoology. University of Aberdeen, Aberdeen AB9
2TN, ScotIand
Diane Stevenson Vertebrate Division, Milwaukee Public Museum, Milwaukee, Wisconsin 53233
Merlin D. Tuttle Vertebrate Division, Milwaukee Public Museum, Milwaukee, Wisconsin 53233
Don E. Wilson U.S. Fish and Wildlife Service. National Museum of Natural
History, Washington, D.C. 20560

vii

Preface
Among living vertebrates bats and birds are unique in their ability to fly, and it is
this common feature that sets them apart ecologically from other groups. Bats are
in some ways the noctumal equivalents of birds, having evolved and radiated
into a diversity of forms to fill many of the same niches. The evolution of flight
and echolocation in bats was undoubtedly a prime mover in the diversification of
feeding and roosting habits, reproductive strategies, and social behaviors. Bats
have successfully colonized almost every continential region on earth (except
Antarctica), as weIl as many oceanic islands and archipelagos. They comprise
the second largest order of mammals (next to rodents) in number of species and
probably exceed all other such groups in overall abundance. Bats exhibit a
dietary diversity (including insects, fruits, leaves, flowers, nectar and pollen,
fish. other vertebrates, and blood) unparalleled among other living mammals.
Their reproductive pattems range from seasonal monestry to polyestry, and
mating systems inelude promiscuity, monogamy, and polygyny.
The vast majority of what we know about the ecology of bats is derived
from studies of only a few of the approximately 850 species, yet in the past two
decades studies on bats have escalated to a level where many important empirical
pattems and processes have been identified. This knowledge has strengthened
our understanding of ecological relationships and encouraged hypothesis testing
rather than perpetuated a catalog of miscellaneous observations. More than ever
before, there is an urgent need to intensify efforts to study the ecology of these
unique mamrnais, both in temperate and tropical regions. The tropical regions of
the world support the highest diversity of bat faunas known, but uniess steps are
taken to decelerate or reverse the current rate of loss of tropical habitats to
altemate land-use pattems, the ecological diversity of this fauna may never be
fully known.
The objective in organizing this volume was to provide a balanced and
authoritative account of major topics on the ecology of bats. Each of the ten
chapters has been prepared by one or more acknowledged experts. The range of
topics treated is by no means exhaustive; rather, the subjects are representative of
ix

Preface

a sizable Iiterature and have not been treated in reeent works. The authors were
eneouraged to develop ehapters in an ecologieal framework based on subjects of
their own interest and expertise, to emphasize empirical studies as well as theory,
and, where appropriate, to inelude discussions of adaptations and evolutionary
trade-offs. It is my hope that this volume will provide points of departure for
more rigorous study by ecologists and ethologists interested in bats and encourage others to consider ecological adaptations of bats as models for comparative
study.
An attempt has been made to organize the sequence of chapters so that most
of them are presented in a logical progression; otherwise they are grouped together by related subjects. An inevitable consequence of a multiauthor volume of
this scope is that it leads to some redundancy in subject matter. Where appropriate, cross-references are made to relevant chapters. As in any discipline, new
viewpoints emerge that will not always be received with universai agreement.
This is as it should be, and consequently I have tried not to interfere with
presentarions of unorthodox ideas and interpretations, for the sake of encouraging further dialogue. The following comments provide an introduction to the
topics included in this volume.
Roosts play a dominant role in the life history of bats, but there have been
few attempts to integrate roosring habits, foraging behavior. social behavior.
morphology, and energetics as factors influencing the roosting ecology of bats.
Taking this approach in preparing Chapter 1, I (Kunz) have summarized the
diversity of roosting adaptations, emphasized factors affecting roost fidelity and
daily time budgets, and provided the first comprehensive treatment of nightroosting ecology. Roost availability. roost dimensions. energetic considerations.
and risks of predation appear to be major determinants of roost use. In recognizing the importance of roosts to successful reproduction. Racey (Chapter 2) draws
extensively from the literature and upon his own work to identify factors controlling reproduction in bats. Environmental factors affecting reproductive cyeles.
rates of embryonic development, and the timing and sequence of births are
examined in bats from both temperate and tropical regions. He emphasizes that
food supply during laetation and weaning is the single most important evolutionary factor in the timing of reproductive cyeles.
Plasticity appears to be one of the IJlost prevaJenr features of growth and
survival in bats. Tuttle and Stevenson (Chaptef 3) draw from an extensive literature and point out that both the litter size and developmental state at birth vary
with biological and environmental factors. They note that successful growth of
pre- and postweaned young varies among species and is markedly affected by
feeding success and roost temperatures. In their anaJyses of bat survival they
caution the reader on the limitations of sampling procedures and statistical treatments in published studies.

Preface

xi

The imponance of body size, food habits, and environmental conditions are
emphasized by McNab's treatment of bat energetics and water balance in Chapter 4. Drawing extensively from his own work on the physiological ecology of
temperate and tropical bats, he discusses factors influencing energy expenditure
in bats, the eeological significanee of energetics (including endotherrny and the
relationship between metabolie rate and life span), energy budgets, and the
evolution ofbat energetics. He argues that the distributionallimits to bats may be
imposed by food habits and an energy eeonomy imposed by small body size.
In Chapter 5 Erken summarizes an extensive literature on aetivity pattems
of bats and, drawing from his own experimental work, interprets the periodicity
of aetivity rhythms as adaptations to environments subjected to a 24-hr rhythm.
He suggests that the flight and foraging activities of different speeies are influenced by the plastieity of a circadian system, the inhibitory aetivity of light, and
the sensitivity of the circadian system to meteorologieal faetors. The central
thesis of Chapter 6, by Findley and Wilson, is that morphologieal traits used by
systematists are valid ecological indices. Findley and Wilson emphasize the
value of multivariate morphologieal analyses for quantifying the ecological
niches of bats and in making comparisons of speeies within and among different
ecological communities.
Fenton (Chapter 7) integrates field studies of bat echolocation with properties of insect prey in the interpretation of the feeding ecology of insectivorous
bats. He suggests that the hearing ability of insects may influence the design of
echolocation calls used by bats when feeding. He notes that most insectivorous
bats are dietary opponunists. Because of extensive overlaps in habitat use and
foraging time and morphological similarities among sympatric species, Fenton
argues that there is no convincing evidence that insectivorous bats compete for
and thus panition available food resources. Foraging strategies of plant-visiting
bats are the subject of Chapter 8, where Aeming examines the imponance of
food availability and roosting behavior to foraging strategies (including foraging
group size and fraging distance). He compares the foraging strategies of four
well-studied species of Neotropical plant-visiting bats and emphasizes the importance of integrating studies on roosting behavior, socialorganization, and feeding behavior to the ultimate understanding of the foraging strategies of bats.
Most mutualistic interactions between plants and planr-visiting bats are
unique to tropical regions, and in Chapter 9, Heithaus synthesizes a diverse
botanicaI literature and integrates it with current knowledge of the foraging
behavior of plant-visiting bats. He develops a model for coupled speciation
between bats and plants. summarizes their complex coadaptations. and discusses
the ecological and evolutionary consequences of bat-plant interactions. He suggests that one should avo id assuming that mutualism between bats and plants is
coevolved, since preadaptations characterize several modem bat-plant systems.

Preface

In spite of the pote!ltial importanee of parasitism in the lives of bats, relatively little research has been published on this topie. Ecologically, the bestknown group of parasites associated with bats is the eetoparasitie inseets, whieh
Marshall treats in Chapter 10. He draws from the literature and from his own
work on life eycIes, host associations, and population dynamies to provide
insight into some rather eomplex host-parasite adaptations unique to these flying
mammals. This ehapter includes an appendix with suggestions for eolleeting
ectoparasitie insects for ecological smdy.
I am grateful to the authors of this volume for enthusiastieally aeeepting the
ehallenge to eontribute their time and ideas and for tolerating my impatienee in
meeting deadlines. The staff of the Plenum Publishing Corporation has been
extremely helpful in all phases of produetion. I owe special thanks to Kirk
Jensen, who initially agreed to embark upon this venture and to Alan Winick and
Daniel Jaul for their indulgenee in my inexperienee as an editor. I want to thank
my most reeent and present graduate smdents, Edythe Anthony, Pete August,
Christopher Bumett, Marty Fujita, Karen Hoying, Al Kurta, and Holly Stack.
for their assistance in proofreading and in the preparation of indices, and Nancy
Tinnel and Christine Zotter, who assisted in elerieal matterso Karl F. Koopman
kindly verified the famiIial and species nomenelature of the Chiroptera. Finally, I
want to acknowledge my graduate mentor, J. Knox Jones. Jr., who encouraged
me to pursue my ecological interests in bats.
Thom"as H. Kunz

Boston. Massachusens

Contents
Chapter 1
Roosting Ecology

Thomas H. Kunz
1. Introduction ............. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
2. Day Roosts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . .. . . . . . . . . . . . . .
2.1. Adaptations for Roosting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
2.2. Roost Activities and Time Budgets . . . . . . . . . . . . . . . . . . . . . ..
2.3. Roost Fidelity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
3. Night Roosts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
3.1. Resting Places. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
3.2. Feeding Perches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
3.3. Feeding Roosts .......................................
3.4. Calling Roosts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
4. Summary ................................................
5. References .............. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..

1
3
3
30
35
37
38
41
42
43
44
46

Chapter 2
Ecology of Bat Reproduction

P. A. Racey
1. Introduction ................ . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
2. The Tirriing of Breeding Seasons . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
2.1. Effeet of Variations in Latitude . . . . . . . . . . . . . . . . . . . . . . . . ..
2.2. Rainfall and Its Effeet on Food Supply. . . . . . . . . . . . . . . . . . ..
3. Environmental Factors Affecting Specific Reproductive Events. . . ..
3.1. Spermatogenesis and Androgenesis . . . . . . . . . . . . . . . . . . . . . ..
3.2. Estrus and Ovulation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
xiii

57
58
59
60
65
65
68

xiv

Contents

3.3. Mating.............................................. 72
3.4. Delayed Fertilization. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 72
~.5. Pregnancyand Lactation ......................... " ..... 75
3.6. Environmental Factors Affecting the Growth and Survival of
young .............................................. 88
3.7. Puberty and Subsequent Fertility and Fecundity . . . . . . . . . . . .. 90
4. Summary ................................................ 92
5. References ....................... . . . . . . . . . . . . . . . . . . . . . . .. 93

Chapter 3
Growth and Survival of Bats
Merlin D. Tuttle and Diane Stevenson
1. Introduction .............................................. 105
2. Prenatal Growth and Development ............................ 106
2.1. Length of Gestation .................................... 106
2.2. Time and Synchrony of Parturition ....................... 107
2.3. Developmental State at Birth ............................ III
2.4. Litter Size ........................................... 113
3. Postnatal Growth and Development ........................... 114
3.1. Preflight ............................................ : 116
3.2. Postflight ............................................ 118
4. Survival ................................................. 120
4.1. Survival Analyses and Results ........................... 1:20
4.2. Survival Determinants .................................. 134
4.3. Survival Strategies ..................................... 138
5. Summary ................................................ 138
6. References ............................................... 139

Chapter 4
Evolutionary Alternatives in the Physiological Ecology of Bats
Brian K. McNab
1. Introduction.............................................. 151
1.1. The Significance of Physiology to the Ecology of Bats . . . . . . . 151
1.2. The Significance of Bats for Physiological Ecology .......... 15:2
2. The Energetics of Bats ...................................... 153
2.1. Factors Determining the Energy Expenditure of Bats ......... 153
2.2. Ecological Significance of Energetics for Bats .............. 163
2.3. Energy Budgets ........... , ........................... 179
2.4. The Evolution of Bat Energetics ......................... 182

Contents

xv

3. The Water Balanee of Bats .................................. 186

3.1. Kidney Function ...................................... 186
3.2. Balancing a Water Budget .............................. 188
4. Distributional Limits to Bats ................................. 189
4.1. Temperate Limits of Tropical Bats ........................ 189
4.2. Limits to Distribution in Temperate Bats ................... 192
5. Summary ................................................ 195
6. References ............................................... 196

Chapter 5
Eeological Aspeets of Bat Aetivity Rhythms
Hans G. Erkert
1. Introduction . . . . . . ........................................ 201
2. Methods for Recording the Activity of Bats ..................... 202
3. Activity Pattems and Timing of Flight Activity under Natural and
Con troll ed Conditions ...................................... 203
3.1. Activity Pattems ...................................... 203
3.2. Arousal and Timing of Flight Activity ..................... 207
3.3. Light-Sampling Behavior ............................... 212
3.4. lnfluence of Extemal Factors on Activity Rhythms .......... 213
4. Activity Rhythms during Hibemation .......................... 220
5. The Endogenous Origin of Bat Activity Rhythms ................ 221
5.1. Circadian Activity Rhythms ............................. 221
5.2. Susceptibility of Period to Exogenous Influences ............ 223
5.3. The Phase Response of Circadian Activity Rhythms
to Light Pulses ........................................ 225
5.4. Entrainment of Circadian Rhythms ....................... 226
5.5. Range of Entrainment and Speed of Resynchronization ....... 229
6. Ecological Adaptation of Circadian Systems and Evolutionary
Aspects .................................................. 233
7. Summary ................................................ 235
8. References ............................................... 236

Chapter 6
Eeological Significance of Chiropteran Morphology
James S. Findley and Don E. Wilson
1. Introduction .............................................. 243
2. The Trophic Niche ......................................... 244
2.1. Flight and Wing Morphology ............................ 244

Contents

3.

4.
S.
6.
7.

2.2. Jaw Morphology and Diet .............................. 247

2.3. Brain Size ........................................... 248
2.4. General Morphology and Feeding ..... : .................. 249
Morphology and Community Strueture ......................... 251
3.1. Speeies Paeking in Temperate versus Tropieal-Bat
Communities ......................................... 252
3.2. Results from Prineipal-Components Analyses ............... 253
Sexual Dimorphism ......................................... 254
Geographie Variation ....................................... 255
Summary ................................................ 256
Referenees '" ............................................ 258

Chapeer 7

Eeboloeation, Inseet Hearing, and Feeding Eeology of Inseetivorous Bats

M. Brock Fenton

1. Introduction .............................................. 261

2. Eeholoeation Calls ......................................... 263
2.1. Call Strueture......................................... 264
2.2. Intensity ............................................. 265
2.3. Frequency ........................................... 266
2.4. Pulse Repetition Rates ................................. 268
2.5. Harmonies ............................................ 269
2.6. Effeetive Range ....................................... 270
3. Hearing and Inseet Defense .................................. 270
4. Responses of Bats to Inseet Hearing ........................... 272
5. Bats as Specialists ......................................... 274
5.1. By Time ............................................. 274
5.2. By Diet ............................................. 274
5.3. By Foraging Strategy .. , ............................... 275
5.4. By Space ............................................ 276
5.5. By Morphology ....................................... 277
5.6. As Rapid Feeders ..................................... 277
6. Other Considerations ....................................... 278
7. Summary ................................................ 280
8. Referenees ............................................... 280

Chapeer 8

Foraging Strategies of Plant-Visiting Bats

Theodore H. Fleming
1. Introduction .............................................. 287

Contents

xvii

2. Food Availability and General Foraging Strategies ............... 290

2.1. Food Availability ...................................... 290
2.2. General Foraging Strategies ............................. 296
3. The Foraging Behavior of Plant-Visiting Bats ................... 302
3.1. Food Habits and Diet Breadth ........................... 303
3.2. Foraging Behavior ..................................... 305
3.3. Case Histories ........................................ 312
4. Summary and General Conclusions ............................ 317
5. References ............................................... 321

Chapter 9
Coevolution between Bats and Plants
E. Raymond Heithaus
1. Introduction .............................................. 327
2. Coupled Speciation ........................................ 328
2.1. Evolutionary Origins of Frugivory and Nectarivory .......... 328
2.2. Effeets of Bats on P!ant Diversification .................... 330
2.3. Coupled Speeiation through Coevolution? ................. 331
3. Complex Coadaptations between Bats and Plants ................ 332
3.1. Coadaptations .................................. : ..... 333
3.2. Aexibility and Diffuse Coevolution ....................... 337
3.3. The Search for Order: Pollination and Dispersal Syndromes ... 343
4. Ecologieal Consequenees of Bat-Plant Interactions ............... 349
4.1. Variation in Effeets .................................... 350
4.2. Demographie Effeets? .................................. 355
4.3. Community Effects .................................... 356
5. Does Coevolution "Matter"? ................................ 358
6. Summary ................................................ 360
7. Referenees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...... 361

Chapter 10
Ecology of Insects Ectoparasitic on Bats
Adrian G. Marshall
1. Introduction .............................................. 369
2. Life Cycles ............................................... 372
2. 1. Pattems ............................................. 372
2.2. Food and Feeding ..................................... 374
2.3. Influenee of Climate and Host Hibernation ................. 374
2.4. Causes of Mortality .................................... 375
2.5. "Number of Generations per Year ...
379
0

xvi

Contents

3. Host Associations .......................................... 379

3.1. Introduction .......................................... 379
3.2. Pattems ............................................. 380
3.3. Reasons ............................................. 382
4. Host Location and Dispersal ................................. 383
4.1. Locomotion .......................................... 383
4.2. Initial Location and Transferenee between Hosts ............ 384
5. Behavior on or Near the Host ................................ 386
5.1. Introduction .......................................... 386
5.2. Pattems ............................................. 386
5.3. Ectoparasites and Host Health ........................... 388
6. Population Dynamies ....................................... 388
6.1. Introduction .......................................... 388
6.2. Pattems and Limits .................................... 389
6.3. Age Structure ......................................... 394
6.4. Sex Ratio ............................................ 394
6.5. Changes in Abundance with Space and Time ............... 395
7. Conclusions .............................................. 395
8. Appendix ................................................ 396
9. References ............................................... 397
Author Index ................................................. 403
Species Index ................................................. 419
Subject Index ................................................. 427