ecology of bats

8/18/2019 Ecology of Bats

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Ecologyo ats

This limited facsimile editian has been issuedfor the purpose of keeping this titl avai ableto the scientific community


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Ecologyo ats

Edited y

Thomas H. KunzBoston UniversityBoston Massachusetts

Plenum Press • New York and London


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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 599.4'045ISBN 13: 978 1 4613 3423 1001: 10.1007/978 1 4613 3421 7

e ISBN 13: 978 1 4613 3421 7

This limited facsimile edition has been issuedfor the purpose of keeping this title availableto the scientific community.

1 9 8 7 6 5 4

© 1982 Plenum Publishing CorporationSoftcover reprint of the hardcover 1 st e ition 1982233 Spring Street, New York, N.Y. 10013

All rights reserved

82-10157

AACR2

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,recording, or otherwise, without written permission from the Publisher


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To Margaret Pamela and David


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ontributors

Hans G Erkert Institut für Biologie III der Universität Tübingen. D-7400Tübingen I Federal Republic of Germany

M. Brock Fenton Department of Biology. Carleton University Ottawa KIS5B6. Canada

James S Findley Department of Biology University of New Mexico. Albuquerque. New Mexico 87 3

Theodore H Fleming Department of Biology. University of Miami CoralGables. 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 GainesvilleFlorida 32611

P. A. Racey Department of Zoology. University of Aberdeen Aberdeen AB92TN ScotIand

Diane Stevenson Vertebrate Division Milwaukee Public Museum Milwaukee Wisconsin 53233

Merlin D. Tutt le Vertebrate Division Milwaukee Public Museum Milwaukee Wisconsin 53233

Don E Wilson U.S. Fish and Wildlife Service. National Museum of NaturalHistory Washington D.C. 20560

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reface

Among living vertebrates bats and birds are unique in their ability to fly, and it isthis common feature that sets them apart ecologically from other groups. Bats arein some ways the noctumal equivalents o birds, having evolved and radiatedinto a diversity o forms to fill many o the same niches. The evolution o flightand echolocation in bats was undoubtedly a prime mover in the diversification ofeeding and roosting habits, reproductive strategies, and social behaviors. Batshave successfully colonized almost every continential region on earth exceptAntarctica), as weIl as many oceanic islands and archipelagos. They comprisethe

second largest ordero

mammals next to rodents)in

numbero

species andprobably exceed all other such groups in overall abundance. Bats exhibit adietary 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, andmating systems inelude promiscuity, monogamy, and polygyny.

The vast majority o what we know about the ecology o bats is derivedfrom studies o only a few o the approximately 850 species, yet in the past twodecades studies on bats have escalated to a level where many important empirical

pattems and processes have been identified. This knowledge has strengthenedour understanding o ecological relationships and encouraged hypothesis testingrather than perpetuated a catalog o miscellaneous observations. More than everbefore, there is n urgent need to intensify efforts to study the ecology o theseunique mamrnais, both in temperate and tropical regions. The tropical regions othe world support the highest diversity o bat faunas known, but uniess steps retaken to decelerate or reverse the current rate o loss o tropical habitats toaltemate land-use pattems, the ecological diversity o this fauna may never befully known.

The objective in organizing this volume was to provide a balanced andauthoritative account o major topics on the ecology o bats. Each o the tenchapters has been prepared by one or more acknowledged experts. The range otopics treated is by no means exhaustive; rather, the subjects are representative o

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Preface x

The imponance of body size, food habits, and environmental conditions areemphasized by McNab s treatment of bat energetics and water balance in Chapter 4. Drawing extensively from his own work on the physiological ecology oftemperate and tropical bats, he discusses factors influencing energy expenditurein bats, the eeological significanee of energetics (including endotherrny and therelationship between metabolie rate and life span), energy budgets, and theevolution ofb t energetics. He argues that the distributionallimits to bats may beimposed by food habits and an energy eeonomy imposed by small body size.

In Chapter 5 Erken summarizes an extensive literature on aetivity pattemsof bats and, drawing from his own experimental work, interprets the periodicityof aetivity rhythms as adaptations to environments subjected to a 24-hr rhythm.

He suggests that the flight and foraging activitiesof

different speeies are influenced by the plastieity of a circadian system, the inhibitory aetivity of light, andthe sensitivity of the circadian system to meteorologieal faetors. The centralthesis of Chapter 6, by Findley and Wilson, is that morphologieal traits used bysystematists are valid ecological indices. Findley and Wilson emphasize thevalue of multivariate morphologieal analyses for quantifying the ecologicalniches of bats and in making comparisons of speeies within and among differentecological communities.

Fenton (Chapter 7 integrates field studies of bat echolocation with proper

tiesof

insect prey in the interpretationof

the feeding ecologyof

insectivorousbats. He suggests that the hearing ability of insects may influence the design ofecholocation calls used by bats when feeding. He notes that most insectivorousbats are dietary opponunists. Because of extensive overlaps in habitat use andforaging time and morphological similarities among sympatric species, Fentonargues that there is no convincing evidence that insectivorous bats compete forand thus panition available food resources. Foraging strategies of plant-visitingbats are the subject of Chapter 8, where Aeming examines the imponance offood availability and roosting behavior to foraging strategies (including foraging

group size and föraging distance). He compares the foraging strategies of fourwell-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 areunique to tropical regions, and in Chapter 9, Heithaus synthesizes a diversebotanicaI literature and integrates it with current knowledge of the foragingbehavior of plant-visiting bats. He develops a model for coupled speciationbetween 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 iscoevolved, since preadaptations characterize several modem bat-plant systems.


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Preface

In spite o the pote ltial importanee o parasitism in the lives o bats, relatively little research has been published on this topie. Ecologically, the bestknown group o parasites associated with bats is the eetoparasitie inseets, whiehMarshall treats in Chapter 10. He draws from the literature and from his ownwork on life eycIes, host associations, and population dynamies to provideinsight into some rather eomplex host-parasite adaptations unique to these flyingmammals. This ehapter includes an appendix with suggestions for eolleetingectoparasitie insects for ecological smdy.

am grateful to the authors o this volume for enthusiastieally aeeepting theehallenge to eontribute their time and ideas and for tolerating my impatienee inmeeting deadlines. The staff o the Plenum Publishing Corporation has beenextremely helpful in all phases o produetion. owe special thanks to KirkJensen, who initially agreed to embark upon this venture and to Alan Winick andDaniel Jaul for their indulgenee in my inexperienee as an editor. want to thankmy 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 o indices, and NancyTinnel and Christine Zotter, who assisted in elerieal matterso Karl F. Koopmankindly verified the famiIial and species nomenelature o the Chiroptera. Finally,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


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Contents

ChapterRoosting Ecology

Thomas H Kunz

1 Introduction 12 Day Roosts 3

2 1 Adaptations for Roosting 32 2 Roost Activities and Time Budgets 302 3 Roost Fidelity 35

3 Night Roosts 373 1 Resting Places 383 2 Feeding Perches 413 3 Feeding Roosts 423 4 Calling Roosts 43

4 Summary 445 References 46

Chapter 2Ecology o at Reproduction

P A Racey

1 Introduction 572 The Tirriing of Breeding Seasons 58

2 1 Effeet of Variations in Latitude 59

2 2 Rainfall and Its Effeet on Food Supply 603 Environmental Factors Affecting Specific Reproductive Events 653 1 Spermatogenesis and Androgenesis 653 2 Estrus and Ovulation 68

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xiv Contents

3.3. M a t i n g 723.4. Delayed Fertilization. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72~ 5 Pregnancyand Lactation 753.6. Environmental Factors Affecting the Growth and Survival of

young 88

3.7. Puberty and Subsequent Fertility and Fecundity . . . . . . . . . . . 904. Summary 925. References . . . . . . . . . . . . . . . . . . . . . . 93

ChapterGrowth and Survival of Bats

Merlin D. Tuttle and Diane Stevenson

1 Introduction 1052. Prenatal Growth and Development 106

2.1. Length of Gestation 1062.2. Time and Synchrony of Parturition 1072.3. Developmental State at Birth2.4. Litter Size 113

3. Postnatal Growth and Development114

3.1. Preflight : 1163.2. Postflight 118

4. Survival 1204.1. Survival Analyses and Results 1:204.2. Survival Determinants 1344.3. Survival Strategies 138

5. Summary 1386. References 139

Chapter 4Evolutionary Alternatives in the Physiological Ecology of Bats

Brian K. McNab

1 I n t r o d u c t i o n 151

1.1. The Significance of Physiology to the Ecology of Bats . . . . . . . 1511.2. The Significance of Bats for Physiological Ecology 15:2

2. The Energetics of Bats 532.1. Factors Determining the Energy Expenditure of Bats 1532.2. Ecological Significance of Energetics for Bats 1632.3. Energy Budgets , 1792.4. The Evolution of Bat Energetics 182


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Contents x

3. The Water Balanee of Bats 1863.1. Kidney Function 1863.2. Balancing a Water Budget 188

4. Distributional Limits to Bats 1894.1. Temperate Limits of Tropical Bats 1894.2. Limits to Distribution in Temperate Bats 192


ChapterEeological Aspeets of Bat Aetivity Rhythms

Hans G. Erkert

1 Introduction . . . . . . 2012 Methods for Recording the Activity o Bats 2023. Activity Pattems and Timing of Flight Activity under Natural and

Con troll ed Conditions 2033.1. Activity Pattems 2033.2. Arousal and Timing o Flight Activity 2073.3. Light Sampling Behavior 2123.4. lnfluence of Extemal Factors on Activity Rhythms 213

4. Activity Rhythms during Hibemation 2205. The Endogenous Origin of Bat Activity Rhythms 221

5.1. Circadian Activity Rhythms 2215.2. Susceptibility of Period to Exogenous Influences 2235.3. The Phase Response of Circadian Activity Rhythms

to Light Pulses 2255.4. Entrainment of Circadian Rhythms 2265.5. Range of Entrainment and Speed of Resynchronization 229

6. Ecological Adaptation of Circadian Systems and EvolutionaryAspects 233


Chapter 6Eeological Significance of Chiropteran Morphology

James S. Findley and Don E Wilson1 Introduction 2432 The Trophic Niche 244

2.1. Flight and Wing Morphology 244


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Contents

2.2. Jaw Morphology and Diet 2472.3. Brain Size 2482.4. General Morphology and Feeding : 249

3. Morphology and Community Strueture 2513.1. Speeies Paeking in Temperate versus Tropieal-Bat

Communities 2523.2. Results rom Prineipal-Components Analyses 253

4. Sexual Dimorphism 254S Geographie Variation 2556. Summary 2567. Referenees ' 258

Chapeer 7Eeboloeation Inseet Hearing and Feeding Eeology of Inseetivorous Bats

M. Brock Fenton

1 Introduction 2612. Eeholoeation Calls 263

2.1. Call Strueture 2642.2. Intensity 265

2.3. Frequency 2662.4. Pulse Repetition Rates 2682 5 Harmonies 2692.6. Effeetive Range 270

3. Hearing and Inseet Defense 2704. Responses of Bats to Inseet Hearing 2725. Bats as Specialists 274

5.1. By Time 2745 2 By Diet 274

5 3 By Foraging Strategy 2755 4 By Space 2765 5 By Morphology 2775 6 As Rapid Feeders 277

6. Other Considerations 2787. Summary 2808. Referenees 280

Chapeer 8Foraging Strategies of Plant-Visiting Bats

Theodore H. Fleming

1. Introduction 287


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Contents xvii

2. Food Availability and General Foraging Strategies 2902.1. Food A vailability 2902.2. General Foraging Strategies 296

3. The Foraging Behavior of Plant-Visiting Bats 3023.1. Food Habits and Diet Breadth 3033.2. Foraging Behavior 3053.3. Case Histories 312

4. Summary and General Conclusions 3175. References 321

Chapter 9Coevolution between Bats and Plants

E. Raymond Heithaus

1. Introduction 3272. Coupled Speciation 328

2.1. Evolutionary Origins of Frugivory and Nectarivory 3282.2. Effeets of Bats on P ant Diversification 3302.3. Coupled Speeiation through Coevolution? 331

3. Complex Coadaptations between Bats and Plants 3323.1. Coadaptations : 3333.2. Aexibility and Diffuse Coevolution 3373.3. The Search for Order: Pollination and Dispersal Syndromes 343

4. Ecologieal Consequenees of Bat-Plant Interactions 3494.1. Variation in Effeets 3504.2. Demographie Effeets? 3554.3. Community Effects 356

5. Does Coevolution Matter ? 3586. Summary 3607. Referenees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361

Chapter 1Ecology of Insects Ectoparasitic on Bats

Adrian G. Marshall

1. Introduction 3692. Life Cycles 372

2. 1. Pattems 3722.2. Food and Feeding 3742.3. Influenee of Climate and Host Hibernation 3742.4. Causes of Mortality 3752.5. Number of Generations per Year 379


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xviü Contents

3 Host Associations 3793 1 Introduction 3793 2 Pattems 3803 3 Reasons 382

4 Host Location and Dispersal 3834 1 Locomotion 3834 2 Initial Location and Transferenee between Hosts 384

5 Behavior on or Near the Host 3865 1 Introduction 3865 2 Pattems 3865 3 Ectoparasites and Host Health 388

6 Population Dynamies 3886 1 Introduction 3886 2 Pattems and Limits 3896 3 Age Structure 3946 4 Sex Ratio 3946 5 Changes in Abundance with Space and Time 395

7 Conclusions 3958 Appendix 3969 References 397

Author Index 403Species Index 419Subject Index 427

ecology of bats

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