Bats of Colima, Mexico 9780806152165, 9780806152387, 0806152389

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Cornelio Sánchez-Hernández ❚ María de Lourdes Romero-Almaraz ❚ Gary D. Schnell Michael L. Kennedy ❚ Troy L. Best ❚ Robert D. Owen ❚ Sara B. González-Pérez

BATS OF COLIMA, MEXICO

Animal Natural History Series Victor H. Hutchison, General Editor

B AT S O F CO L I M A ,

ME XICO Cornelio Sánchez-Hernández María de Lourdes Romero-Almaraz Gary D. Schnell Michael L. Kennedy Troy L. Best Robert D. Owen Sara B. González-Pérez

University of Oklahoma Press : Norman In cooperation with the Sam Noble Oklahoma Museum of Natural History

This book is made possible by the generous support of the Sam Noble Oklahoma Museum of Natural History.

Illustrations on title page spread: (left to right) northern ghost bat, photo by Gary D. Schnell; Wagner’s mustached bat, photo by Cornelio Sánchez-Hernández (see fig. 6.66, p. 143); western yellow bat, photo by Silvia S. Zalapa (see fig. 6.114, p. 200); Waterhouse’s leaf-nosed bat, photo by María de Lourdes Romero-Almaraz; big free-tailed bat, photo by Silvia S. Zalapa (see fig. 6.95, p. 177); and hoary bat, photo by Silvia S. Zalapa (see fig. 6.110, p. 196).

Library of Congress Cataloging-in-Publication Data Names: Sánchez Hernández, Cornelio. Title: Bats of Colima, Mexico / Cornelio Sánchez-Hernández [and six others]. Description: Norman : University of Oklahoma Press, 2016. | Series: Animal natural history series ; volume 14 | “In cooperation with the Sam Noble Oklahoma Museum of Natural History.” | Includes bibliographical references and index. Identifiers: LCCN 2016001067 | ISBN 978-0-8061-5216-5 (pbk. : alk. paper) Subjects: LCSH: Bats—Mexico—Colima (State) Classification: LCC QL737.C5 S328 2016 | DDC 599.40972/36—dc23 LC record available at https://lccn.loc.gov/2016001067 Bats of Colima, Mexico is Volume 14 in the Animal Natural History Series. The paper in this book meets the guidelines for permanence and durability of the Committee on Production Guidelines for Book Longevity of the Council on Library Resources, Inc. ∞ Copyright © 2016 by the University of Oklahoma Press, Norman, Publishing Division of the University. Manufactured in the U.S.A. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise—except as permitted under Section 107 or 108 of the United States Copyright Act—without the prior written permission of the University of Oklahoma Press. To request permission to reproduce selections from this book, write to Permissions, University of Oklahoma Press, 2800 Venture Drive, Norman, OK 73069, or email [email protected]. 1 2 3 4 5 6 7 8 9 10

To the people of Colima. Their hospitality, kindness, and cooperation over many years were crucial to the successful completion of this book and fostered many pleasant lifelong memories.

CONTENTS List of Illustrations . . . . . . . List of Tables . . . . . . . . . Preface . . . . . . . . . . . Acknowledgments . . . . . . . List of Acronyms for Museums with   Bat Specimens from Colima . . .

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. ix . xii . xiii . xv

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1. INTRODUCTION TO STUDY OF COLIMA BATS . 3 2. STUDY AREA . . . . . . . . . . . . . . 9 Physiography . . . . . . . . . . . . . . . 10 Geology . . . . . . . . . . . . . . . . . 11 Hydrology . . . . . . . . . . . . . . . . 12 Climate . . . . . . . . . . . . . . . . . 12 Vegetation . . . . . . . . . . . . . . . . 12 Mangrove . . . . . . . . . . . . . . . 15 Halophytic vegetation . . . . . . . . . . . 15 Tropical deciduous forest . . . . . . . . . . 16 Tropical semideciduous forest . . . . . . . . 16 Oak forest . . . . . . . . . . . . . . . 17 Pine-oak forest . . . . . . . . . . . . . 17 Secondary vegetation . . . . . . . . . . . 18 Agricultural land . . . . . . . . . . . . . 18

3. METHODS . . . . . . . . . . . . . . . 21 Field Studies . . . . . . . . . . . . . . . 21 Mist nets . . . . . . . . . . . . . . . . 22 Roosts . . . . . . . . . . . . . . . . . 22 Collection and preservation of specimens . . . . 22 Identification and Location of Specimens . . . . . 22 Measurements . . . . . . . . . . . . . . 23 Accounts . . . . . . . . . . . . . . . . 24 Description, distribution, and conservation status . 24 Environmental information . . . . . . . . . 24 Diet . . . . . . . . . . . . . . . . . 25 Reproduction . . . . . . . . . . . . . . 25 Activity and other observations . . . . . . . 26 Measurements, specimens examined, localities,   and records in literature . . . . . . . . . 27 Gazetteer . . . . . . . . . . . . . . . . 27

4. ANALYSIS . . . . . . . . . . . . . . . . 29 Specimens in Museums and Documented in Literature 29 Natural History . . . . . . . . . . . . . . 35 Diet . . . . . . . . . . . . . . . . . 35 Reproductive pattern . . . . . . . . . . . 36 Zoogeographic Associations . . . . . . . . . 38 Conservation and Endemic Species . . . . . . . 38 Misidentified Species from Colima . . . . . . . 38

5. KEY TO BATS OF COLIMA . . . . . . . . . 41 Families . . . . . . . . . . . . . . . . . 43 Genera and Species of Emballonuridae . . . . . . 46 Subfamilies of Phyllostomidae . . . . . . . . . 46 Genera and Species of Glossophaginae . . . . . . 47 Genera and Species of Phyllostominae . . . . . . 49 Genera and Species of Stenodermatinae . . . . . 49 Genera and Species of Mormoopidae . . . . . . 51 Species of Natalidae . . . . . . . . . . . . . 51 Genera and Species of Molossidae . . . . . . . . 51 Genera and Species of Vespertilionidae . . . . . . 53

6. ACCOUNTS . . . . . . . . . . . . . . . 57 FAMILY EMBALLONURIDAE (Sac-winged Bats) . 57 Gray sac-winged bat, Balantiopteryx plicata . . . 58 Northern ghost bat, Diclidurus albus . . . . . . 61 Greater sac-winged bat, Saccopteryx bilineata . . . 63 FAMILY PHYLLOSTOMIDAE (Leaf-nosed Bats) . . 66 Subfamily Desmodontinae Common vampire bat, Desmodus rotundus . . . 67 Subfamily Glossophaginae Geoffroy’s tailless bat, Anoura geoffroyi . . . . . 70 Mexican long-tongued bat, Choeronycteris mexicana 73 Commissaris’s long-tongued bat,   Glossophaga commissarisi . . . . . . . . . 75 Gray’s long-tongued bat, Glossophaga leachii . . . 77 Western long-tongued bat, Glossophaga morenoi . 79 Pallas’s long-tongued bat, Glossophaga soricina . . 81 Mexican long-nosed bat, Leptonycteris nivalis . . 84 Lesser long-nosed bat, Leptonycteris yerbabuenae . 87 Banana bat, Musonycteris harrisoni . . . . . . 90 Subfamily Phyllostominae Tricolored big-eared bat, Glyphonycteris sylvestris . 93 Waterhouse’s leaf-nosed bat, Macrotus waterhousii 95 Common big-eared bat, Micronycteris microtis . . 98

vii

Subfamily Carolliinae Gray short-tailed bat, Carollia subrufa . . . . . 101 Subfamily Stenodermatinae Honduran yellow-shouldered bat,   Sturnira hondurensis . . . . . . . . . . 103 Northern yellow-shouldered bat, Sturnira parvidens 106 Intermediate fruit-eating bat, Artibeus intermedius 109 Jamaican fruit-eating bat, Artibeus jamaicensis . . 112 Great fruit-eating bat, Artibeus lituratus . . . . 116 Wrinkle-faced bat, Centurio senex . . . . . . . 119 Salvin’s big-eyed bat, Chiroderma salvini . . . . 121 Aztec fruit-eating bat, Dermanura azteca . . . . 123 Pygmy fruit-eating bat, Dermanura phaeotis . . . 125 Toltec fruit-eating bat, Dermanura tolteca . . . . 128 Velvety fruit-eating bat, Enchisthenes hartii . . . 131 FAMILY MORMOOPIDAE (Mustached Bats) . . . 133 Peters’s ghost-faced bat, Mormoops megalophylla . 134 Davy’s naked-backed bat, Pteronotus davyi . . . 137 Common mustached bat, Pteronotus parnellii . . 140 Wagner’s mustached bat, Pteronotus personatus . . 143 FAMILY NOCTILIONIDAE (Bulldog Bats) . . . . 146 Greater bulldog bat, Noctilio leporinus . . . . . 147 FAMILY NATALIDAE (Funnel-eared Bats) . . . . 150 Woolly funnel-eared bat, Natalus lanatus . . . . 151 Mexican greater funnel-eared bat,   Natalus mexicanus . . . . . . . . . . . 153 FAMILY MOLOSSIDAE (Free-tailed Bats) . . . . 156 Mexican dog-faced bat, Cynomops mexicanus . . 157 Fierce bonneted bat, Eumops ferox . . . . . . . 159 Underwood’s bonneted bat, Eumops underwoodi . 161 Pallas’s mastiff bat, Molossus molossus . . . . . 164 Black mastiff bat, Molossus rufus . . . . . . . 166

Sinaloan mastiff bat, Molossus sinaloae . . . . . 168 Peale’s free-tailed bat, Nyctinomops aurispinosus . 171 Pocketed free-tailed bat, Nyctinomops femorosaccus 173 Broad-eared free-tailed bat,   Nyctinomops laticaudatus . . . . . . . . . 175 Big free-tailed bat, Nyctinomops macrotis . . . . 177 Big crested mastiff bat, Promops centralis . . . . 180 Brazilian free-tailed bat, Tadarida brasiliensis . . 182 FAMILY VESPERTILIONIDAE (Common Bats) . . 185 Subfamily Vespertilioninae Brazilian brown bat, Eptesicus brasiliensis . . . . 186 Argentinian brown bat, Eptesicus furinalis . . . . 188 Big brown bat, Eptesicus fuscus . . . . . . . . 190 Western red bat, Lasiurus blossevillii . . . . . . 193 Hoary bat, Lasiurus cinereus . . . . . . . . . 196 Northern yellow bat, Lasiurus intermedius . . . . 198 Western yellow bat, Lasiurus xanthinus . . . . . 200 Little yellow bat, Rhogeessa parvula . . . . . . 202 Mexican big-eared bat, Corynorhinus mexicanus . 205 Townsend’s big-eared bat,   Corynorhinus townsendii . . . . . . . . . 207 Subfamily Myotinae Silver-tipped myotis, Myotis albescens . . . . . 210 Southwestern myotis, Myotis auriculus . . . . . 212 Californian myotis, Myotis californicus . . . . . 214 Carter’s black myotis, Myotis carteri . . . . . . 216 Cinnamon myotis, Myotis fortidens . . . . . . 218 Dark-nosed small-footed myotis, Myotis  melanorhinus . . . . . . . . . . . . . 220 Fringed myotis, Myotis thysanodes . . . . . . 223 Cave myotis, Myotis velifer . . . . . . . . . 225 Yuma myotis, Myotis yumanensis . . . . . . . 228

Appendix A. Gazetteer . . . . . . . . . . . . 231 Appendix B. Measurements . . . . . . . . . . . 243 Appendix C. Skulls . . . . . . . . . . . . . . 259 Glossary . . . . . . . . . . . . . . . . . 277 Literature Cited . . . . . . . . . . . . . . . 285 Illustration Credits . . . . . . . . . . . . . . 307 Index . . . . . . . . . . . . . . . . . . . 309

viii

CONTENTS

ILLUSTRATIONS

1.0

Volcán de Fuego, 2008–2009 .

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1.1 Field crew members for expeditions in 1996–97 and 2008–2009 . . . . . . . . . . 1.2

Field camp during 2008–2009 expedition .

1.3

Schnell, Kennedy, and Lara, 2006

1.4

Meneses near La Yerbabuena, 2008 .

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5.0 Sánchez-Hernández extricating intermediate fruit-eating bat from mist net, 2006 . . . . . . . 40

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5.2 Dorsal, ventral, and lateral views of cranium and lateral view of mandible . . . . .

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2.0 Ruiz-Gutiérrez and Akins with bats in irrigation tunnel, 2006 . . . . . . . . . . . . . . . 8 2.1

Map: localities of bats recorded in Colima .

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2.2

Map: elevation gradients in Colima .

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10

2.3

Map: slopes in Colima . . . . . . . . . . . 11

2.4

Map: mean annual precipitation in Colima . . . . . 13

2.5

Map: mean annual temperature in Colima . . . . . 13

2.6

Map: five basic climate regions in Colima .

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2.7 Map: locations of vegetation types, agricultural lands, and urban areas . . . . . . . . . . . . . 14 2.8

Mangrove in dry season . . . . . . . . . . . 15

2.9

Halophytic vegetation .

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15

2.10 Tropical deciduous forest in dry and wet season .

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2.11 Tropical semideciduous forest .

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2.12 Oak forest . . . . . . . . . . . . . . . 17 2.13 Pine-oak forest .

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42

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5.3

Dentition of generalized bat .

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Basic components of skull of young bat . . . . . . 43

5.5 Characteristics: noseleaf, tail vertebrae, W-shaped molar, divided upper lip, wing sac . . . . . . . . . . 44 5.6 Characteristics: postorbital process, tail, bonnet-shaped ears, natalid organ, keratin capsule, rostrum with hexagonal depression . . . . . . . 45 5.7 Characteristics: long thumb, rostrum shape, uropatagium condition, incisor configurations . . . . . . . . 47 5.8 Characteristics: incisor configurations, pterygoid wings, ear shape, wing pattern . . . . . .

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. 48

5.9 Characteristics: nasal bones absent, molar configuration, uropatagium border, hairs on feet . . . . . . . . 50 5.10 Characteristics: lip, incisor configuration, molar shape, uropatagium fringe . . . . . . . . 52 6.0 Intermediate fruit-eating bat (Artibeus intermedius) caught in mist net . . . . . . . . . . . 6.1 Family Emballonuridae: gray sac-winged bats (Balantiopteryx plicata) . . . . . . .

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2.14 Secondary vegetation .

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6.2

Gray sac-winged bat

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6.3

Map: known distribution of gray sac-winged bat in Colima

6.4

Northern ghost bat (Diclidurus albus)

6.5

Map: known distribution of northern ghost bat in Colima . 62

6.6

Greater sac-winged bat (Saccopteryx bilineata) . . . . 63

2.15 Agricultural lands

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External anatomy of a bat . . . . . . . . . . 41

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3.0 Students Sánchez Vázquez, Alcántara, Amman, and Salas-Rojas, 2001 . . . . . . . .

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20

3.1

Coleaders of the 1999–2000 expedition . . . . . . 21

3.2

Owen in 2001 and González-Pérez in 2007 . . . . . 21

3.3

Young of lesser long-nosed bat (Leptonycteris yerbabuenae)

4.0 Poindexter exhibiting Peters’s ghost-faced bat, 2007 . 4.1

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26 . 28

Number of specimens collected, by decade . . . . . 29

4.2 Number of specimens of bats from Colima in North American collections . . . . . .

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4.3 Number of species of bats from Colima in North American collections . . . . .

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4.4 Cumulative number of species of bats in museums and literature, by year . . .

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4.5 Number of species and museum specimens per family for bats in Colima . . . . . . . . . 35

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6.9

6.11 Geoffroy’s tailless bat (Anoura geoffroyi)

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6.12 Map: known distribution of Geoffroy’s tailless bat . . . 71 6.13 Mexican long-tongued bat (Choeronycteris mexicana) .

. 73

6.14 Map: known distribution of Mexican long-tongued bat in Colima . . . . . . . . . . . . . .

. 74

6.15 Pregnant Commissaris’s long-tongued bat (Glossophaga commissarisi) . . . . . . . . . . 75

Reproductive patterns of bats in Colima .

6.17 Gray’s long-tongued bat (Glossophaga leachii)

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. 57 .

6.10 Map: known distribution of common vampire bat in Colima 69

4.7

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Common vampire bat (Desmodus rotundus) . . . . . 67

6.16 Map: known distribution of Commissaris’s long-tongued bat in Colima . . . . .

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6.8 Family Phyllostomidae: lesser long-nosed bats (Leptonycteris yerbabuenae) . . . . . .

Diets of bats in Colima . . . . . . . . . . . 36 .

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6.7 Map: known distribution of greater sac-winged bat in Colima . . . . . . . . . . . . .

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ix

6.18 Map: known distribution of Gray’s long-tongued bat in Colima . . . . . . . . . . . . .

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. 78

6.57 Velvety fruit-eating bat (Enchisthenes hartii) . . . .

6.19 Western long-tongued bat (Glossophaga morenoi) . . . 79

6.58 Map: known distribution of velvety fruit-eating bat in Colima . . . . . . . . . . . . .

6.20 Map: known distribution of western long-tongued bat in Colima . . . . . . . . . . . . . .

6.59 Family Mormoopidae: colony in cave (Pteronotus parnellii and P. personatus)

6.21 Pallas’s long-tongued bat (Glossophaga soricina)

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6.60 Peters’s ghost-faced bat (Mormoops megalophylla) . . . 134

6.22 Map: known distribution of Pallas’s long-tongued bat in Colima . . . . . . . . . . . . . . . 83

6.61 Map: known distribution of Peters’s ghost-faced bat in Colima . . . . . . . . . . . . .

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. 135

6.23 Mexican long-nosed bat (Leptonycteris nivalis)

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6.62 Davy’s naked-backed bat (Pteronotus davyi) . . . .

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6.24 Map: known distribution of Mexican long-nosed bat in Colima . . . . . . . . . . . . .

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6.63 Map: known distribution of Davy’s naked-backed bat in Colima . . . . . . . . . . . . .

6.25 Lesser long-nosed bat (Leptonycteris yerbabuenae) .

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. 87

6.64 Common mustached bat (Pteronotus parnellii) .

6.26 Map: known distribution of lesser long-nosed bat in Colima

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6.27 Banana bat (Musonycteris harrisoni) . . . . . . . 90

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6.65 Map: known distribution of common mustached bat in Colima . . . . . . . . . . . . . 6.66 Wagner’s mustached bat (Pteronotus personatus) .

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. 143

6.28 Map: known distribution of banana bat in Colima .

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6.29 Tricolored big-eared bat (Glyphonycteris sylvestris) .

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6.67 Map: known distribution of Wagner’s mustached bat in Colima . . . . . . . . . . . . .

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. 144

. 94

6.68 Family Noctilionidae: greater bulldog bat (Noctilio leporinus) fishing over pond . . . . . .

146

6.30 Map: known distribution of tricolored big-eared bat in Colima . . . . . . . . . . . . .

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6.31 Waterhouse’s leaf-nosed bat (Macrotus waterhousii) .

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6.32 Map: known distribution of Waterhouse’s leaf-nosed bat in Colima . . . . . . . . . . . . . . 6.33 Common big-eared bat (Micronycteris microtis)

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. 95 . 96

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6.34 Map: known distribution of common big-eared bat in Colima 99 6.35 Gray short-tailed bat (Carollia subrufa) . . . . . . 101 6.36 Map: known distribution of gray short-tailed bat in Colima

102

6.37 Honduran yellow-shouldered bat (Sturnira hondurensis) . 103 6.38 Map: known distribution of Honduran yellow-shouldered bat in Colima . . . . . . . . . . . . . . 104 6.39 Northern yellow-shouldered bat (Sturnira parvidens) . . 106 6.40 Map: known distribution of northern yellow-shouldered bat in Colima . . . . . . . . . . . . . 6.41 Intermediate fruit-eating bat (Artibeus intermedius)

. 108

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6.42 Map: known distribution of intermediate fruit-eating bat in Colima . . . . . . . . . . . .

. 109 .

. 110

6.43 Jamaican fruit-eating bat (Artibeus jamaicensis) . . .

112

6.44 Map: known distribution of Jamaican fruit-eating bat in Colima . . . . . . . . . . . . . .

. 114

6.45 Great fruit-eating bat (Artibeus lituratus) . . . . .

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6.46 Map: known distribution of great fruit-eating bat in Colima 117 6.47 Wrinkle-faced bat (Centurio senex) .

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6.48 Map: known distribution of wrinkle-faced bat in Colima . 120 6.49 Salvin’s big-eyed bat (Chiroderma salvini) .

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6.50 Map: known distribution of Salvin’s big-eyed bat in Colima

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6.51 Aztec fruit-eating bat (Dermanura azteca)

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6.52 Map: known distribution of Aztec fruit-eating bat in Colima . . . . . . . . . . . . 6.53 Pygmy fruit-eating bat (Dermanura phaeotis) .

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6.54 Map: known distribution of pygmy fruit-eating bat in Colima . . . . . . . . . . . . .

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6.55 Toltec fruit-eating bat (Dermanura tolteca) . . . . . 128 6.56 Map: known distribution of Toltec fruit-eating bat in Colima . . . . . . . . . . . . .

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ILLUSTRATIONS

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. 129

6.69 Greater bulldog bat . . . . . . . . . . . . 147 6.70 Map: known distribution of greater bulldog bat in Colima 148 6.71 Family Natalidae: maternity colony, Mexican greater funnel-eared bat (Natalus mexicanus) . . . . . . 6.72 Woolly funnel-eared bat (Natalus lanatus)

150

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6.73 Map: known distribution of woolly funnel-eared bat in Colima . . . . . . . . . . . . .

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6.74 Mexican greater funnel-eared bat (Natalus mexicanus)

. 152 .

153

6.75 Map: known distribution of Mexican greater funnel-eared bat in Colima . . . . . . . . . . . . . . 154 6.76 Family Molossidae: pocketed free-tailed bat (Nyctinomops femorosaccus) . . . . .

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6.77 Mexican dog-faced bat (Cynomops mexicanus) .

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6.78 Map: known distribution of Mexican dog-faced bat 6.79 Fierce bonneted bat (Eumops ferox)

. 156 .

. 157 . 158

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6.80 Map: known distribution of fierce bonneted bat 6.81 Underwood’s bonneted bat (Eumops underwoodi)

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. 161

6.82 Map: known distribution of Underwood’s bonneted bat in Colima . . . . . . . . . . . . . . 6.83 Pallas’s mastiff bat (Molossus molossus) .

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. 162 . 164

6.84 Map: known distribution of Pallas’s mastiff bat in Colima 6.85 Black mastiff bat (Molossus rufus) .

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165 . 166

6.86 Map: known distribution of black mastiff bat in Colima . 167 6.87 Sinaloan mastiff bat (Molossus sinaloae) . . . . . . 168 6.88 Map: known distribution of Sinaloan mastiff bat in Colima

169

6.89 Peale’s free-tailed bat (Nyctinomops aurispinosus) . . . 171 6.90 Map: known distribution of Peale’s free-tailed bat in Colima 172 6.91 Pocketed free-tailed bat (Nyctinomops femorosaccus) . . 173 6.92 Map: known distribution of pocketed free-tailed bat in Colima . . . . . . . . . . . . .

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. 174

6.93 Broad-eared free-tailed bat (Nyctinomops laticaudatus) .

175

6.94 Map: known distribution of broad-eared free-tailed bat in Colima . . . . . . . . . . . . . .

. 176

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225

6.96 Map: known distribution of big free-tailed bat in Colima . 178

6.137 Map: known distribution of cave myotis in Colima . .

226

6.97 Big crested mastiff bat (Promops centralis) .

6.138 Yuma myotis (Myotis yumanensis)

228

6.95 Big free-tailed bat (Nyctinomops macrotis) .

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6.98 Map: known distribution of big crested mastiff bat in Colima . . . . . . . . . . . . .

. 180 .

. 181

6.99 Brazilian free-tailed bat (Tadarida brasiliensis) . . . . 182 6.100 Map: known distribution of Brazilian free-tailed bat in Colima . . . . . . . . . . . . .

.

. 184

6.101 Family Vespertilionidae: northern yellow bat (Lasiurus intermedius) . . . . . . . . . . . 185 6.102 Brazilian brown bat (Eptesicus brasiliensis) . . . . . 186 6.103 Map: known distribution of Brazilian brown bat in Colima 187 6.104 Argentinian brown bat (Eptesicus furinalis) . . . . . 188 6.105 Map: known distribution of Argentinian brown bat in Colima . . . . . . . . . . . . . 6.106 Big brown bat (Eptesicus fuscus) .

.

.

.

.

.

. .

. 189 . 190

6.107 Map: known distribution of big brown bat in Colima . . 191 6.108 Western red bat (Lasiurus blossevillii) . . . . . .

193

6.109 Map: known distribution of western red bat in Colima .

194

6.110 Hoary bat (Lasiurus cinereus) . . . . . . . . . 196 6.111 Map: known distribution of hoary bat in Colima . 6.112 Northern yellow bat (Lasiurus intermedius) .

.

. .

. 197 .

. 198

6.113 Map: known distribution of northern yellow bat in Colima 199 6.114 Western yellow bat (Lasiurus xanthinus) .

.

.

.

.

6.115 Map: known distribution of western yellow bat in Colima

. 200 201

6.116 Little yellow bat (Rhogeessa parvula) . . . . . . . 202 6.117 Map: known distribution of little yellow bat in Colima .

203

6.118 Mexican big-eared bat (Corynorhinus mexicanus) . . . 205 6.119 Map: known distribution of Mexican big-eared bat in Colima . . . . . . . . . . . . .

.

. 206

6.120 Townsend’s big-eared bat (Corynorhinus townsendii) . . 207 6.121 Map: known distribution of Townsend’s big-eared bat in Colima . . . . . . . . . . . . . . 6.122 Silver-tipped myotis (Myotis albescens)

. 208

. . . . . . 210

6.123 Map: known distribution of silver-tipped myotis in Colima . . . . . . . . . . . .

.

.

. 211

6.124 Southwestern myotis (Myotis auriculus) . . . . . . 212 6.125 Map: known distribution of southwestern myotis in Colima . . . . . . . . . . . .

.

.

. 213

6.136 Cave myotis (Myotis velifer)

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

6.139 Map: known distribution of Yuma myotis in Colima . . 229 . . . . . 230

A.0

Map: localities of bats recorded in Colima

A.1

Schnell and Kennedy at Playa de Oro, 2005

.

.

.

.

. 231

C.1 Skulls: Balantiopteryx plicata, Diclidurus albus, Saccopteryx bilineata, Desmodus rotundus . . . . . . . . . 260 C.2 Skulls: Anoura geoffroyi, Choeronycteris mexicana, Glossophaga commissarisi, G. leachii . . . . . . . 261 C.3 Skulls: Glossophaga morenoi, G. soricina, Leptonycteris nivalis, L. yerbabuenae . . . . . . . 262 C.4 Skulls: Musonycteris harrisoni, Glyphonycteris sylvestris, Macrotus waterhousii, Micronycteris microtis . . . .

263

C.5 Skulls: Carollia subrufa, Sturnira parvidens, S. hondurensis, Artibeus intermedius . . . . . . . 264 C.6 Skulls: Artibeus jamaicensis, A. lituratus, Centurio senex, Chiroderma salvini . .

.

.

. 265

C.7 Skulls: Dermanura azteca, D. phaeotis, D. tolteca, Enchisthenes hartii . . . . . . . . . .

.

.

.

. 266

C.8 Skulls: Mormoops megalophylla, Pteronotus davyi, P. parnellii, P. personatus . . . . . . . . . .

267

C.9 Skulls: Noctilio leporinus, Natalus lanatus, N. mexicanus, Cynomops mexicanus . . . . . . . . . . . 268 C.10 Skulls: Eumops ferox, E. underwoodi, Molossus molossus, M. rufus . . . . . . . . . . . . . . . 269 C.11 Skulls: Molossus sinaloae, Nyctinomops aurispinosus, N. femorosaccus, N. laticaudatus . . . . . . . . 270 C.12 Skulls: Nyctinomops macrotis, Promops centralis, Tadarida brasiliensis, Eptesicus brasiliensis . . . . . 271 C.13 Skulls: Eptesicus furinalis, E. fuscus, Lasiurus blossevillii, L. cinereus . . . . . . . . . . . . . .

272

C.14 Skulls: Lasiurus intermedius, L. xanthinus, Rhogeessa parvula, Corynorhinus mexicanus . . . .

273

C.15 Skulls: Corynorhinus townsendii, Myotis albescens, M. auriculus, M. californicus . . . . . . .

.

. 274

C.16 Skulls: Myotis carteri, M. fortidens, M. melanorhinus, M. thysanodes . . . . . . . . . . . . .

275

C.17 Skulls: Myotis velifer, M. yumanensis . . . . . . . 276

6.126 Californian myotis (Myotis californicus) . . . . . . 214 6.127 Map: known distribution of Californian myotis in Colima 215 6.128 Carter’s black myotis (Myotis carteri) .

.

.

.

.

.

. 216

6.129 Map: known distribution of Carter’s black myotis in Colima . . . . . . . . . . . .

.

.

. 217

6.130 Cinnamon myotis (Myotis fortidens) . . . . . . . 218 6.131 Map: known distribution of cinnamon myotis in Colima . 219 6.132 Dark-nosed small-footed myotis (Myotis melanorhinus) . 220 6.133 Map: known distribution of dark-nosed small-footed myotis in Colima . . . . . . . . . . . . 6.134 Fringed myotis (Myotis thysanodes)

6.135 Map: known distribution of fringed myotis in Colima .



221

. . . . . . . 223

G.0 Sánchez-Hernández checking for Davy’s naked-backed bats, 2007 . . . . .

.

.

.

.

.

277

LC.0 Headquarters for several annual expeditions, Playa de Oro, 2000 . . . . . . . . . . . . 284 CR.0 Sánchez-Hernández, Lara Ortiz, and González-Pérez with northern yellow-shouldered bat . . . . . I.0

.

. 306

.

. 308

ILLUSTRATIONS

xi

Pacific sunset at Rancho Majagua .

.

.

.

.

.

. 224

TABLES

1.1

Species of bats known from Colima . . . . . . . . 7 .

.

.

.

.

.

.

4.1

Species of bats recorded in Colima

4.2

Diets of bats in Colima . . . . . . . . . . . 37

4.3

Reproductive patterns of bats in Colima .

.

.

.

.

32 . 39

B.1 Five external measurements and mass of bats from Colima . . . . . . . . . . . . . . 243 B.2 Two wing and three skull measurements of bats from Colima . . . . . . . . . . . . . . 249 B.3

xii

Five skull measurements of bats from Colima

. . . . 254

PREFACE The American coauthors of this book first developed an interest in the fauna of Colima, Mexico, in the early 1970s, during visits to the Hacienda El Cóbano, a facility owned by the University of Oklahoma and located in the small village of El Cóbano, near the city of Colima. Gary Schnell was a new assistant professor in the Department of Zoology, and curator of birds at what was then the Stovall Museum of Science and History at the University of Oklahoma. Gary’s predecessor as curator of birds was the distinguished emeritus professor George Miksch Sutton, an internationally known ornithologist and artist who had conducted pioneering fieldwork studying birds in Mexico as early as the 1930s, and who was still very active. Friends of George who were interested in birds knew of his experience and expertise on the subject of Mexican birds, and knew as well about the range of outreach programs at Hacienda El Cóbano being rapidly developed by the University of Oklahoma through an office overseen by Richard Handcock. George’s friends urged him to lead a bird-watching trip to Colima; George was interested but reluctant to take on sole responsibility for leading the trip. He suggested that Gary be asked to be a coleader, and Gary readily accepted. The initial trip to Colima went well and further trips watching and studying birds quickly followed, some involving the general public, some for students, and some a combination of the two. Gary’s formal training was in ornithology, but he had an interest in mammals as well. Among his first assemblage of graduate students were Mike Kennedy and Troy Best. Both had primary interest and considerable expertise in mammals, and their interest was shared by Robert Owen, then an undergraduate. Gary urged them to participate in the Colima trips, and it really did not take much urging, given that they were already conducting some mammal studies in Mexico. Gary struck a very informal deal with these students: everyone go to Colima, and when the mammal-oriented students were not studying birds, they were free to work with mammals. The study of mammals in Colima by the Oklahoma stem group was on. The group even had some specimens of mammals from Colima for reference in the Stovall Museum of the University of Oklahoma, given that the director of the museum at that time, Keever Greer, had done some collecting in the state and had briefly visited the Hacienda. The early work by the Oklahoma group led to the collection of numerous bats, mostly from in and around El Cóbano

but occasionally from more distant locales. The early trips involved numerous undergraduate and graduate students (and occasionally faculty), many adventures and challenges, and a fondness and respect for Mexico and Mexican culture that persist today. Additionally, during these early trips to Colima, the research group met numerous people and made many friends. From the start, two individuals (Esteban Meneses and Angel Lara) contributed significantly to its success in Colima. Angel was the director of the Hacienda El Cóbano, and Esteban was a young businessman in Colima City; both became lifelong friends and go-to people if the group needed something or had a problem. It would be a mistake to omit mention of Mary Lara, who contributed significantly in keeping Hacienda El Cóbano running smoothly during our visits. In recognition of Esteban’s significant contributions in fostering scientific study of the fauna of Colima, he was presented a Meritorious Service Award by the Southwestern Association of Naturalists in 2006. In the mid-1970s (following graduation), Troy went to Northeastern University in Boston, Mike went to the University of Memphis, and Robert finished his undergraduate degree at the University of Oklahoma and started a PhD program with Gary at Oklahoma. During the late 1970s, Mike took trips to Colima from Memphis and was joined on numerous occasions by Troy. Mick Harvey (faculty member and bat expert at Memphis) accompanied Mike and Troy on most of these trips. In 1984, Mike, Troy, and Mick published a paper dealing with the bats of Colima. The breadth of the group working in Colima was growing, along with the scholarship. The 1970s were glory days for travel and work in Colima for the stem group from Oklahoma, but in the 1980s circumstances changed. During a difficult time financially, the University of Oklahoma sold the Hacienda El Cóbano. At Memphis, travel and other expenses made it problematic to work in Mexico. The result was that no one in the group went to Colima during the 1980s. However, this did not dampen the group’s enthusiasm and interest in study in this part of Mexico. Members of this informal group often gathered at the annual meeting of the Southwestern Association of Naturalists, and talk invariably turned to events that occurred during the early Colima trips, and the strong desire to go back. One memorable experience that often surfaced during these discussions was the Campo Uno sojourn, a collecting trip in a remote mountainous area and one characterized as

xiii

a true misadventure. There were significant vehicle damages due to a flooded road, amateur road-building, and numerous other surprises. As the group neared the village of Campo Uno with their final destination in sight, the late afternoon sun indicated they needed to turn around and head back down the mountain. There was simply no time to explore, and they all speculated about what they had missed. A common statement thereafter was that the group needed to go back to Campo Uno. This continued for numerous annual meetings and over numerous beers, but eventually members of the group had had enough. It was decided that they needed to do one thing or the other—actually plan and make a trip to Colima, or stop talking about it. The challenge was on. As a result, in December 1996 Gary Schnell and the others initiated the first of what turned out to be fourteen annual expeditions to study the mammalian fauna of Colima. Each trip started on December 30th and ended on January 14th (from 1996–97 through 2009–2010). The mammalian fauna in Colima previously had received little scientific attention, even though Colima is situated in a rich biodiversity hotspot on the western coast of Mexico. This second round of study in Colima began with a phone call from Mike Kennedy to Gary Schnell in the spring of 1996. They agreed on the need to focus on mammals in Colima and also recognized that times had changed; it was both necessary and beneficial to involve Mexican mammalogists in these studies. At the time, Robert Owen was working with a group from Mexico studying mammals in Michoacán, a state adjacent to Colima, and he knew that some of his colleagues in Michoacán would be interested in working on this research. An international team was coming together and eventually included Schnell, Owen, Kennedy, and Best from the United States and Cornelio Sánchez-Hernández and María de Lourdes (Lulú) Romero-Almaraz of the Universidad Nacional Autónoma de México in Mexico City. In addition, Sara B. González-Pérez, an undergraduate working with Cornelio and Lulú, accompanied the group on several early trips and later completed a master’s degree with Gary at the University of Oklahoma. She was a solid contributor on the later trips as well and, throughout, served as a living alarm clock, singing everyone awake almost every morning in the field. Two other faculty participants from Mexico on the initial trip were Ricardo López-Wilchis of the Universidad Autónoma Metropolitana, Unidad Iztapalapa in Mexico City, and Arturo Núñez-Garduño of the Universidad Michoacana de San Nicolás de Hidalgo at Morelia, Michoacán. Not surprisingly, the first collecting site during this series of trips was Campo Uno and the surrounding cloud-forest habitat. This time, the United States group made it. It remains one of their treasured collecting sites in Colima. In addition to Cornelio and Lulú’s active participation in the field and their bringing a host of student participants each xiv

PREFACE

year, they took on major and vitally important responsibilities over the course of these studies. Cornelio played a critical role in obtaining the needed permits for conducting studies in Colima, as well as in the post-trip processing of specimens (e.g., cleaning skulls). Esteban Meneses and Cornelio often had the responsibility of communicating with landowners and others to obtain permission to work in various parts of the state. Throughout the many years of this collaboration, Lulú, with help from various students, took on the crucial task of continually updating a detailed database of all specimens collected. Furthermore, Cornelio and Lulú made a major commitment to the overall project by visiting numerous museums in Mexico and the United States to examine Colima specimens, and by spending many days measuring specimens. These have all been important tasks that helped hold the overall enterprise together and made possible a comprehensive assessment of the bat fauna of Colima. From the beginning, the annual trips involved faculty and students (both graduate and undergraduate) from several universities in Mexico and the United States. They have involved well over three hundred students and twenty or more faculty members from various institutions in Mexico and the United States. Collectively, there were from sixteen to thirty-six participants each year. Students came from many types of backgrounds but with a commonality of adventure and an anticipation of hands-on learning. Students had the opportunity to receive academic credit if they desired; however, most students elected to participate just for the experience. For students, the trips were lessons in geography, interpersonal relations, field biology, and (sometimes) extending outside their personal comfort zone. Most trips cemented memories, and many lifetime friendships developed within and between participants from the two countries. Stories from those trips have been told over and over again. Several students participated in trips to Colima and then on later trips when they held faculty or other professional positions. For faculty, rewards were similar to those garnered by students. Many rank their experiences in Colima as among the most rewarding of their careers. As teachers, they are grateful to have had the opportunity to bring students from two countries together to learn from each other and to learn about biology in a country and state so rich in biodiversity. There are many treasures in Colima—its people, art, history, and culture. However, a treasure often overlooked is its biodiversity, including the multitude of species of both plants and animals. The authors hope that their studies of bats contribute in a small way to the sustainability of this treasured biodiversity, foster an interest in biodiversity for others to follow, and contribute toward collaboration of future ecological research involving biologists and colleagues in Mexico and the United States.

ACKNOWLEDGMENTS Given that our research endeavors in Colima have spanned more than four decades, there are many individuals who have been valued participants in this enterprise and deserve thanks. Faculty and other professionals beyond the authors who have participated in one or more field expeditions to Colima for mammal studies include Mel L. Beck, H. Delano Black, Frederick F. B. Elder, Joseph A. Grzybowski, Michael J. Harvey, Ricardo López-Wilchis, Arturo Núñez-Garduño, Howard H. Thomas, Bill A. Simco, and Earl G. Zimmerman. In addition to this list, five individuals initially went to Colima with us as students and then again after becoming faculty members or taking some other type of professional position: José Antonio Guerrero, Cheryl A. Goudie, Phyllis K. Kennedy, Juliann L. Waits, and Michael C. Wooten. As indicated in the preface, more than three hundred students have accompanied us to Colima, a list too long to cite here. With respect to the fourteen annual trips, there were students who were involved in a major way, taking part in three or more trips. These included Luz Eugenia Alcántara Quintana, Adolfo Calderón Zarza, Gilberto Federico García Ruiz, Ezequiel Guerrero Ibarra, Charles H. Kilgore, Laura Angélica Lara Ortiz, Sara Thelma Martínez Chapital, Anacaren Morales Ortiz, Cassie J. Poindexter, Mónica Salas-Rojas, Leobardo Sánchez Vázquez, and Alejandro Taboada Salgado. We received special assistance from a number of individuals. Paola Ivette Orozco Nolasco, César Emiliano Escalona Prado, and Jenifer Ruth Segura Ramos helped in developing the glossary for this book. Mitzi Atzel Reyes Romero assisted in preparation of skulls and with the creation of composite photographs of selected skulls for inclusion in appendix C. Laura Angélica Lara Ortiz and Anacaren Morales Ortiz provided notable assistance in entering, updating, and proofreading data contributed to the mammal database for Colima. Gerardo Arévalo G., Miguel Ángel Martínez Vilchis, and Georgina Ortega Leite of the library at the Instituto de Biología, Universidad Nacional Autónoma de México, facilitated our work by obtaining numerous electronic copies of relevant publications. Albino Luna Sánchez provided excellent drawings used in conjunction with the dichotomous

keys. After Sara González-Pérez created the initial environmental maps and several of us worked to generate locality maps, Carol Zuber-Mallison added her creative touches in designing the final map products. We also thank those who have provided photographs for use in this book (see illustration credits). Don E. Wilson carefully read the manuscript and provided valuable comments and suggestions. Ana “Kiki” E. Hiott and Victoria L. Smyth were critical contributors relative to our being able to field the fourteen annual expeditions; they assisted in generating funds that were used to provide equipment and support for these trips. Additional financial support was provided by the Coordinación de la Investigación Científica through the Instituto de Biología of the Universidad Nacional Autónoma de México—to facilitate extended international visits during preparation of the manuscript—as well as by the Sam Noble Oklahoma Museum of Natural History. Appropriate permits were obtained from the Instituto Nacional de Ecología, Dirección General de Vida Silvestre FAUT.103 to Cornelio Sánchez-Hernández. Research protocols were approved by the Animal Care and Use Committee of the University of Oklahoma. We are appreciative of the numerous landowners in Colima who gave us permission to conduct fieldwork on their lands, which was essential for our being able to sample adequately the numerous habitats in the state. These landowners included Abelardo Ahumada, Trinidad Alcaraz, Rubén Anaya, Jesús Ballesteros Silva, Javier Contreras, Francisco del Toro López, José Guadalupe Valle Pérez, Porfirio Gutierrez, Felipe Guzmán, Seferino Huezo, José Magaña Sandoval, Antonio Pérez de la Torre, Gilberto Rocha Contreras, and Jorge Vizcaíno González. As indicated in the preface, Esteban Meneses and Angel Lara, residents of Colima, provided valuable logistic and other assistance to our groups over many years. Without their assistance, our accomplishments in Colima would have been very much diminished. Finally, we appreciate the fact that our families have been tolerant of our attraction to Colima and of our many days (and New Year’s Eves) away from home as we pursued our fascination with the bats that live in Colima.

xv

ACRONYMS FOR MUSEUMS WITH BAT SPECIMENS FROM COLIMA AMNH BMUNLV

xvi

American Museum of Natural History (New York, New York) University of Nevada, Las Vegas (Las Vegas)

CNMA

Colección Nacional de Mamíferos, Instituto de Biología, Universidad Nacional Autónoma de México (Mexico, D.F.)

FMNH

Mammal Collection, Field Museum of Natural History (Chicago, Illinois)

IPN

Colección de Mamíferos, Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional (Mexico, D.F.)

KU

University of Kansas Natural History Museum (Lawrence)

LACM

Natural History Museum of Los Angeles County (Los Angeles, California)

LSUMZ

Louisiana State University Museum of Natural Science (Baton Rouge)

MSB

Museum of Southwestern Biology, University of New Mexico (Albuquerque)

MVZ

Museum of Vertebrate Zoology, University of California at Berkeley (Berkeley)

NMNH

United States National Museum of Natural History (Washington, D.C.)

OMNH

Mammal Collection, Sam Noble Oklahoma Museum of Natural History, University of Oklahoma (Norman)

ROM

Royal Ontario Museum (Toronto, Ontario, Canada)

TTU

Museum of Texas Tech University (Lubbock)

UAEM

Colección de Mamíferos, Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Morelos (Cuernavaca)

UAM-I

Universidad Autónoma Metropolitana, Unidad Iztapalapa (Mexico, D.F.)

UAZ

Mammal Collection, University of Arizona (Tucson)

UM

Mammal Collection, University of Memphis (Memphis, Tennessee)

UMMZ

University of Michigan Museum of Zoology (Ann Arbor)

WNMU

Mammal Collection, Western New Mexico University (Silver City)

BATS OF COLIMA, MEXICO

Figure 1.0. View of the active Volcán de Fuego that greeted expedition members in 2008– 2009 when they emerged from their tents.

1

INTRODUCTION TO STUDY OF COLIMA BATS

The state of Colima is in the transition zone between two major biogeographic regions of the Americas, the Nearctic and the Neotropical. As a result, both temperate and tropical species inhabit the region. Highly variable topography, climate, and vegetation—along with the influence of ocean currents—favor creation of numerous ecological niches that are occupied by many species of organisms. Therefore, Colima has a high diversity of mammals, and it is also within the area having the largest number of endemic species in Mexico (Ramírez-Pulido and Müdespacher 1987; Fa and Morales 1993; Ceballos and Oliva 2005; Ceballos 2014). For Colima, 66 species of bats have been documented, which represents 47.5% of the 138 species recognized for Mexico by Ramírez-Pulido et al. (2014) plus the intermediate fruit-eating bat (Artibeus intermedius). Of those species in Colima, 5 (7.6%) are endemic to Mexico (Simmons 2005). This high diversity is remarkable considering that the surface area of Colima is only 5,191 km2 (INEGI 2014). For comparison, Michoacán, an adjacent state, has a surface area of 58,585 km2—more than ten times the area of Colima—and supports 75 species of bats (Núñez 2005). Jalisco, the other state adjacent to Colima, has a surface area of 80,386 km2 and 73 bat species (Godinez et al. 2011). Colima is included within a region considered to be among the three most diverse hotspots for biodiversity in the world (Myers et al. 2000). This biodiversity hotspot is second in diversity of vertebrates and endemic species of vertebrates, as well as second in diversity of mammals and first in number of endemic mammals (Myers et al. 2000). Given this rich faunal heritage, it is of both heuristic and practical importance to have a comprehensive inventory of species of bats that inhabit the state, and information about their natural history. Early investigators provided information on the bat fauna of Colima. Audley C. Buller traveled and collected extensively in Mexico, and in 1890 the American Museum of Natural History (AMNH) received two specimens of Davy’s naked-backed bat (Pteronotus davyi) from Colima taken 15 January 1890 on the “Plains of Colima.” The first major effort to collect mammals specifically in Colima was by Edward W. Nelson and Edward A. Goldman, who arrived at the Port of Manzanillo, Colima, on 24 January 1892 (Young 1947). As a result of their three months of fieldwork, more than 245 specimens representing 17 species of bats from Colima were deposited in the National Museum of Natural History (NMNH). Most specimens were from Manzanillo, Hacienda La Magdalena, and the city of Colima. A few specimens from Colima from that trip are housed in the Field Museum of Natural History (FMNH). In 1950, J. R. Alcorn, from the University of Kansas, visited the Manzanillo area and collected six species of bats; another collecting trip followed from the University of Kansas in 1961 by M. R. Lee. The American Museum also received specimens of bats taken in Colima in 1956, when J. A. Davis, E. Gould, and W. J. Schaldach, Jr., collected specimens near Pueblo Juárez and Manzanillo. Beginning in early 1958, significant efforts to collect bats in Colima were made by personnel associated with the Natural History Museum of Los Angeles County (LACM). These expeditions occurred from 1958 to 1970 and again in 1975; collectors were A. L. Gardner, W. J. Schaldach, Jr., P. L. Clifton, D. R. Patten, P. A. Flanagan, and Ciro González B., a Mexican naturalist collector from Colima. As a result, 2,452 specimens of 53 species of bats were collected. The Natural History Museum of Los Angeles County has the only specimens available from Colima for nine of those species, as well as the type specimen of the banana bat (Musonycteris harrisoni). In 1965, 3

Ciro González B. also collected 143 bats of 24 species for the Louisiana State University Museum of Natural Science (LSUMZ). The first bat specimen deposited in a Mexican collection—the Colección Nacional de Mamíferos (CNMA)—was a Wagner’s mustached bat (Pteronotus personatus) collected by I. Sanderson in 1941. E. Gray and R. Boolootina collected and deposited additional bat specimens in that collection in 1954. Bernardo Villa R. and Ciro González B. collected specimens of bats near Pueblo Juárez and El Mixcoate in 1955, 1961, and 1962. Other specimens were collected over the years, but those seem to have resulted more from occasional opportunistic fieldwork than focused research. The Mammal Collection of the Instituto Politéctico Nacional sent Aurelio Ocaña to collect in Colima in 1965, J. Nava in 1970, and M. Aguilar C. in 1988.

Figure 1.1. Above: Members of the field crew for the 1996–97 expedition. Below: Members of the field crew for the 2008– 2009 expedition, shown under a large fig (Ficus) tree.

4

BATS OF COLIMA, MEXICO

Figure 1.2. Field camp during the 2008–2009 expedition.

Figure 1.3. Left to right: Gary Schnell, Mike Kennedy, and Angel Lara in 2006 discuss multiple trips made to Colima in the 1970s, when expeditions were headquartered at the Hacienda El Cóbano. Angel served as the Director of the Hacienda.

In the preface, we briefly outlined the collecting efforts in the 1970s by personnel associated with the University of Oklahoma and later the University of Memphis, Auburn University, and Texas Tech University. During that period, 1,258 specimens were collected, 551 deposited in what is now the Sam Noble Oklahoma Museum of Natural History (OMNH) of the University of Oklahoma, and 707 at the University of Memphis; those at Memphis were later transferred to the museum in Oklahoma. Furthermore, the fourteen joint field expeditions (figs. 1.1–1.4) outlined in the preface (1996–97 through 2009–2010) yielded 3,394 specimens of bats. In the 1960s, the literature began to include more detailed information on bats occurring in Colima. Villa-R. (1966) noted 37 species of bats from Colima as part of an inventory of bats in Mexico. Kennedy et al. (1984) published a list of bats mainly from the regions of Playa de Oro and

INTRODUCTION TO THE STUDY OF BATS

5

Figure 1.4. Esteban Meneses in 2008 near La Yerbabuena. Throughout our work, and particularly in the 1990s and 2000s, Esteban served as an indispensable colleague in Colima, making arrangements for access to field and camping sites, providing advice on where to obtain needed services, and helping establish contact with relevant government officials and other individuals in Colima.

El Cóbano. Sánchez-Hernández et al. (2002) supplied new records for 13 species of bats in Colima and provided additional information for 15 other species. In addition, Sánchez-Hernández et al. (2009) documented 18 species of bats from the Ranchitos ejido. Other publications recognized one or two species of bats from within the state (Gardner 1962a, 1962b, 1963, 1966; Best and Kennedy 1984). The total number of known species in Colima now stands at 66 (table 1.1). Studies in Colima have also provided information on flight of a number of species. Flight speeds have been evaluated for the following: gray sac-winged bats (Balantiopteryx plicata) (Kennedy et al. 1977); Peters’s ghost-faced bats (Mormoops megalophylla), Davy’s naked-backed bats (Pteronotus davyi), common mustached bats (P. parnellii), and Wagner’s mustached bats (P. personatus; Kennedy et al. 1977; Hopkins et al. 2003); common vampire bats (Desmodus rotundus; Sánchez-Hernández et al. 2006); Pallas’s long-tongued bats (Glossophaga soricina), gray short-tailed bats (Carollia subrufa), and Mexican greater funnel-eared bats (Natalus mexicanus, referenced as N. stramineus; Akins et al. 2007). Overall, in spite of the relatively large number of species and specimens collected in Colima, the chiropteran fauna has received little attention in the literature from mammalogists. Until now, neither a monograph nor a comprehensive book has been produced concerning the bats of Colima. Having a comprehensive list of bats in a region, and knowledge of their distributions within that region, are important starting points for many kinds of basic and applied research, such as that aimed at understanding ecological processes in areas with similar or varied environmental conditions. Regarding conservation concerns, this type of information is relevant for decision makers who designate protected natural areas and specific conservation measures for threatened or rare species. Given the opportunities we have had over several decades to study bats in Colima, coupled with the rich source of information available through museum specimens collected in the state, we have undertaken the following: (1) to compile a list of the species of bats occurring in the state of Colima; (2) to summarize a wide range of information concerning each species; (3) to develop an identification key to species based on external, cranial, and dental characteristics; (4) to report on geographic distribution of each species in Colima; and (5) where possible, to characterize environmental conditions where species occur in the state, as well as where they have not been documented. 6

BATS OF COLIMA, MEXICO

Table 1.1. SPECIES OF BATS KNOWN FROM COLIMA FAMILY EMBALLONURIDAE Balantiopteryx plicata (gray sac-winged bat) Diclidurus albus (northern ghost bat) Saccopteryx bilineata (greater sac-winged bat) FAMILY PHYLLOSTOMIDAE Subfamily Desmodontinae Desmodus rotundus (common vampire bat) Subfamily Glossophaginae Anoura geoffroyi (Geoffroy’s tailless bat) Choeronycteris mexicana (Mexican long-tongued bat) Glossophaga commissarisi (Commissaris’s long-tongued bat) Glossophaga leachii (Gray’s long-tongued bat) Glossophaga morenoi (western long-tongued bat) Glossophaga soricina (Pallas’s long-tongued bat) Leptonycteris nivalis (Mexican long-nosed bat) Leptonycteris yerbabuenae (lesser long-nosed bat) Musonycteris harrisoni (banana bat) Subfamily Phyllostominae Glyphonycteris sylvestris (tricolored big-eared bat) Macrotus waterhousii (Waterhouse’s leaf-nosed bat) Micronycteris microtis (common big-eared bat) Subfamily Carolliinae Carollia subrufa (gray short-tailed bat) Subfamily Stenodermatinae Sturnira hondurensis (Honduran yellow-shouldered bat) Sturnira parvidens (northern yellow-shouldered bat) Artibeus intermedius (intermediate fruit-eating bat) Artibeus jamaicensis (Jamaican fruit-eating bat) Artibeus lituratus (great fruit-eating bat) Centurio senex (wrinkle-faced bat) Chiroderma salvini (Salvin’s big-eyed bat) Dermanura azteca (Aztec fruit-eating bat) Dermanura phaeotis (pygmy fruit-eating bat) Dermanura tolteca (Toltec fruit-eating bat) Enchisthenes hartii (velvety fruit-eating bat) FAMILY MORMOOPIDAE Mormoops megalophylla (Peters’s ghost-faced bat) Pteronotus davyi (Davy’s naked-backed bat) Pteronotus parnellii (common mustached bat) Pteronotus personatus (Wagner’s mustached bat)



FAMILY NOCTILIONIDAE Noctilio leporinus (greater bulldog bat) FAMILY NATALIDAE Natalus lanatus (woolly funnel-eared bat) Natalus mexicanus (Mexican greater funnel-eared bat) FAMILY MOLOSSIDAE Cynomops mexicanus (Mexican dog-faced bat) Eumops ferox (fierce bonneted bat) Eumops underwoodi (Underwood’s bonneted bat) Molossus molossus (Pallas’s mastiff bat) Molossus rufus (black mastiff bat) Molossus sinaloae (Sinaloan mastiff bat) Nyctinomops aurispinosus (Peale’s free-tailed bat) Nyctinomops femorosaccus (pocketed free-tailed bat) Nyctinomops laticaudatus (broad-eared free-tailed bat) Nyctinomops macrotis (big free-tailed bat) Promops centralis (big crested mastiff bat) Tadarida brasiliensis (Brazilian free-tailed bat) FAMILY VESPERTILIONIDAE Subfamily Vespertilioninae Eptesicus brasiliensis (Brazilian brown bat) Eptesicus furinalis (Argentinian brown bat) Eptesicus fuscus (big brown bat) Lasiurus blossevillii (western red bat) Lasiurus cinereus (hoary bat) Lasiurus intermedius (northern yellow bat) Lasiurus xanthinus (western yellow bat) Rhogeessa parvula (little yellow bat) Corynorhinus mexicanus (Mexican big-eared bat) Corynorhinus townsendii (Townsend’s big-eared bat) Subfamily Myotinae Myotis albescens (silver-tipped myotis) Myotis auriculus (southwestern myotis) Myotis californicus (Californian myotis) Myotis carteri (Carter’s black myotis) Myotis fortidens (cinnamon myotis) Myotis melanorhinus (dark-nosed small-footed myotis) Myotis thysanodes (fringed myotis) Myotis velifer (cave myotis) Myotis yumanensis (Yuma myotis)

INTRODUCTION TO THE STUDY OF BATS

7

Figure 2.0. In 2006, Fernando Ruiz-Gutiérrez (front) and James Akins (back) share an irrigation tunnel with Mexican greater funnel-eared bats (Natalus mexicanus) and Pallas’s long-tongued bats (Glossophaga soricina).

2

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Colima is one of the smallest states in Mexico, with only 0.3% of the surface area of the country (INEGI 2014). It is in west-central Mexico, bordered by Jalisco on the west, north, and east; Michoacán to the east and south; and the Pacific Ocean to the west and south. It has a surface area of 5,191 km2, the coastline extends 157 km, and it is not far from the equator, with a latitude from 19°31’ to 18°41’N. Its longitude is from 103°29’ to 104°41’W (INEGI 2014). The name “Colima” is from the Nahuatl collimaitl, with colli meaning “ancestors or gods” and maitl indicating “domain of.” The state also includes the Revillagigedo Islands, which consist of four volcanic islands named Benito Juárez, Clarión, San Benedicto, and Roca Partida. No bat has been recorded on any of these islands. On the mainland, Colima is partitioned into ten municipalities, each named after its principal city or town: Armería, Colima, Comala, Coquimatlán, Cuauhtémoc, Ixtlahuacán, Manzanillo, Minatitlán, Tecomán, and Villa de Álvarez (fig. 2.1). The largest cities in the state are Colima (the capital), Comala, Cuauhtémoc, Manzanillo, Tecomán, and Armería. Herein, we use Colima to refer to the state and “city of Colima” to refer to the capital. Figure 2.1. Map of Colima Pac

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18 45 37 9 10 8 19 20 21 22 11 13 38 Minatitlán 23 COMALA 14 27 40 41 24 15 12 25 42 44 3945 43 29 17 28 Cuauhtémoc 46 16 30 5253 47 31 50 Comala 33 49 VILLA DE 48 32 147 34 51 MINATITLÁN 55 56 122 35 36 ÁLVAREZ Villa de 148 26 57 54 58 59 Álvarez 149 60 123 151 CUAUHTÉMOC 124 66 68 70 84 125 Colima 153 83 61 62 154 65 126 150 COQUIMATLÁN 72 82 152 86 63 64 67 85 157 73 69 128 71 138 127 159 155 129 87 76 77 156 89 137 74 75 81 133 135 88 Coquimatlán 158 130 MANZANILLO 90 160 80 91 92 132 94 93 COLIMA 140139 143 161 134 95 96 102 106 107 108 144 142 136 162 99 105 166 109 141 145 131 97 101 110 163 176 167 169 98 100 104 164 165 168 Manzanillo 103 114 177 146 170 113 111 112 197 171 115 199 218 ARMERÍA 117 198 116 201 219 179 118 200 178 180 119 220 221 222 120 181 202 182 223 Ixtlahuacán 172 183 203 186 185 173 187188 174 IXTLAHUACÁN 184 204 121 189 205 190 Armería 175 224 191 194 Tecomán 206 225 192 195 226 193 207 TECOMÁN 227 196 209 208 210 211 213

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Localities of Bats Recorded in Colima

showing all localities for bats. Numbers correspond to those in the gazetteer (appendix A). Dashed lines indicate boundaries of municipalities, which are named after a city or town in each municipality. Principal cities and towns are shown (shaded), as are bodies of water (outlined). Inset map in upper left indicates location of the state of Colima in Mexico. Copyright © 2016, University of Oklahoma Press.

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Figure 2.2. Elevation gradients JALISCO

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Colima lies at the interface of the Nearctic and Neotropical biogeographic regions. Within the state, the Nearctic region includes the highlands of the Volcán de Fuego and the Sierra de Manantlán in northern Colima. The Neotropical region includes slopes and lowlands throughout the state.

Physiography Colima, with its highly variable topography, is part of two physiographic provinces, with 83.7% in the Sierra Madre del Sur and 16.3% in the Eje Volcánico Transversal (INEGI 2014). A transect through Colima, drawn along a line from the summit of Volcán de Fuego (3,820 m) to sea level at the mouth of the Armería River near Boca de Pascuales, is steep and rugged. Few other locales along the Pacific coast of Mexico present such a steep elevational gradient: the actual straight-line distance of 56 km is comparatively short (Schaldach 1963), as shown in figure 2.2. Steep slopes are characteristic of many parts of Colima (fig. 2.3). The massive twin volcanoes—Volcán de Fuego (3,820 m), one of the most active volcanoes in the world, and Volcán de Nieve (4,260 m) in Jalisco—dominate the northern border of Colima and are an integral and vital part of the region. Due to a paucity of roads and trails, the southern slopes of the volcanoes, in Colima, remain relatively inaccessible. However, on the northern slopes, in Jalisco, there are access roads into the dense forests of the taller peak, the Volcán de Nieve. All of the north-northeastern slopes and most of the eastern slopes of these volcanoes are in Jalisco (Schaldach 1963). Beyond coastal ranges, the generally level but much dissected central peneplain (the Plains of Colima) begins (Schaldach 1963). It has an average elevational range from 360 to 1,200 m, with 10

BATS OF COLIMA, MEXICO

Figure 2.3. Slopes in Colima. Pac ifi c

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COLIMA COMALA

Minatitlán

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(in degrees)

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localities such as the city of Colima, Junta Municipal Madrid, Tecolapa, Cuauhtémoc, Trapiche, Puerta de Anzar, Tepames, and Palmillas. The plain tilts gently upward to the north to meet the base of the volcanoes (with communities such as La Yerbabuena, Laguna La María, and San Antonio). Rising to the north are large mountain systems wholly within Colima. These include La Media Luna, Cerro Grande, Cerro Chino, Cerro del Otate, and Cerro del Zacate. These mountains are nearly uniform in maximum height, ranging from 1,830 to 2,130 m.

Geology Geologically, the Quaternary and Tertiary periods of the Cenozoic era and the Cretaceous period of the Mesozoic era are represented in Colima (INEGI 2014). The Quaternary is represented along the coast, excluding the mountains that rise near Playa de Oro and northwest of the bays of Manzanillo and Santiago, as well as Punta El Carrizal. The Quaternary is also exemplified in a narrow central area and in the east-central part of the state. Evidence of the Tertiary can be seen mainly in northern and eastern Colima, as well as near Sierra Perote and Cerro El Pelón. Elevation in this area ranges from 500 m to greater than 1,500 m and includes most of central, northeastern, eastern, and southeastern Colima. The Cretaceous area includes most mountainous regions of northern, northwestern, central, and southeastern Colima. In addition, this geological period is evident in the higher mountains, such as Cerro Grande, Jumpiche, Barrigón, San Diego, Del Toro, La Mina, San Miguel, San Gabriel, and La Yerbabuena. In southern Colima, the Cretaceous period is represented by several mountains 500–700 m in elevation, such as those north of the Cuyutlán and Armería lagoons, and west of Rincón de López.



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Hydrology There are two hydrologic regions in Colima: the Costa de Jalisco region that includes the Chacala-Purificación and Cihuatlán river basins in western Colima, with a surface area of 1,624 km2; and the Armería-Coahuayana region, formed by the Armería and Coahuayana river basins, which together cover the remaining 3,567 km2 of the state (INEGI 2014). The Armería River is the major river system that divides the state along a general north-south line and flows into the Pacific Ocean at Boca de Pascuales. Because of the large size of the river, it has an extensive flood plain. The Coahuayana and El Naranjo rivers form the eastern border of the state. It is the more northerly El Naranjo River that joins the Barreras River from Michoacán to form the Coahuayana River. The Salado River originates at the southeastern slope of Volcán de Fuego and flows southeasterly across the state to join the Coahuayana River. The Cihuatlán River (on some maps the Marabasco River) constitutes part of the western border of Colima. Both the relatively high annual precipitation on the Sierra de Manantlán and the springs on the volcanoes are important sources of drinking water and irrigation for communities in nearby valleys (Schaldach 1963; INEGI 2014).

Climate The higher slopes of the volcanoes are cold throughout the year, with snow often mantling Jalisco’s Volcán de Nieve (also a prominent visual feature in Colima) in January and February. Winter frosts are locally common in the highest valleys (i.e., those with an elevation >2,300 m). Other places with a similar climate are Cerro Grande and Campo Uno. Lowlands and lower mountains of Colima have a typical western Mexican tropical climate. Highly seasonal, Colima is characterized by a pronounced wet season from mid-June to October that alternates with an equally pronounced dry season from November through the middle of June. The dry season is notable for its long succession of clear, cloudless, and warm days. Beginning about mid-June, brief but torrential rains fall almost daily, with rivers and streams flooding in September. Humidity reaches an average of 85% and temperatures average 32°C. Occasional cyclonic tropical storms strike the coast of Colima (Schaldach 1963). Figure 2.4 shows mean annual precipitation that varies from about 600 mm in the broad valleys to 1,700 mm in cloud forests at the highest elevations (WorldClim data). Figure 2.5 shows mean annual temperatures, which range from 9.6°C to 27.0°C (WorldClim data). Based on 1980 environmental data from the Instituto Nacional de Estadística y Geografía (INEGI 1980), five climate types are represented in Colima based on the climate-classification system of Wladimir Köppen as modified for Mexico by García (1964). As shown in figure 2.6, the five types are as follows: (1) semidry very warm with annual mean temperature >22°C in the warmest month and >18°C in the coldest month, with precipitation-to-temperature ratio >22.9; (2) tropical subhumid with annual mean temperature >22°C and summer rainfall; (3) semiwarm subhumid with annual mean temperature 18–22°C and summer rainfall; (4) temperate subhumid with annual mean temperature 12–18°C and summer rainfall; and (5) semicold subhumid with mean annual temperature from 5 to >12°C and summer rainfall. The latter characterizes a small northern portion of Colima close to the peak of the Volcán de Fuego at elevations greater than 2,500 m.

Vegetation Varied topography, diverse geology, a highly seasonal climate combined with moist westerlies from the Pacific Ocean, and the combination of temperate and tropical elements all contribute to the complex mosaic of vegetation that exists in Colima (Rzedowski 1978; Ferrusquía-Villafranca 1998). Cover types for land, based on surface area and the classification system we employed (see below) include secondary vegetation, 51.5%; agricultural lands, 32.7%; forests, 10.9%; mangrove and halophytic vegetation, 1.8%; urban, 1.6%; and bodies of water, 1.5%.

12

BATS OF COLIMA, MEXICO

Figure 2.4. Mean annual Pac

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COLIMA

Coquimatlán

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Precipitation

Ixtlahuacán IXTLAHUACÁN

(in millimeters)

575–600 >600–750 >750–900 >900–1,050 >1,050–1,200 >1,200–1,500 >1,500–1,682

18°45'

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Armería

Tecomán TECOMÁN

MICHOACÁN

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Figure 2.5. Mean annual Pac

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VILLA DE Comala ÁLVAREZ Villa de Álvarez Colima COQUIMATLÁN

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Manzanillo ARMERÍA 19°00'

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Ixtlahuacán IXTLAHUACÁN

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9.6–16.0 >16.0–18.0 >18.0–20.0 >20.0–22.0 >22.0–24.0 >24.0–26.0 >26.0–27.0 104°30'

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Figure 2.6. Five basic climate regions in Colima. Copyright © 2016, University of Oklahoma Press.

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Climate

Ixtlahuacán IXTLAHUACÁN

Semidry very warm Tropical subhumid Semiwarm subhumid Temperate subhumid Semicold subhumid

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Figure 2.7. Locations of six types of natural vegetation in Colima in addition to secondary vegetation, agricultural lands, and urban areas.

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Minatitlán

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COQUIMATLÁN MANZANILLO

Comala Villa de Álvarez Colima

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Vegetation Mangrove Halophytic vegetation Tropical deciduous forest Tropical semideciduous forest Oak forest Pine-oak forest Secondary vegetation Agricultural land 104°30'

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The tree flora of Colima includes about 550 species, with many of these belonging to the families Magnoliaceae, Fabaceae, Euphorbiaceae, and Rubiaceae. Genera with the greatest number of species are Quercus (oaks), Bursera (burseras), Ficus (wild figs), Lonchocarpus (lancepods), and Senna (sennas). The tropical deciduous forest contains 67.1% of the species, while the tropical semideciduous forest includes 33.5% (Padilla-Velarde et al. 2006). Based on land use and vegetation data from INEGI (2011), six types of natural vegetation occur in Colima (fig. 2.7); we are designating these as mangrove, halophytic vegetation, tropical deciduous forest, tropical semideciduous forest, oak forest, and pine-oak forest. Delimiting their extent is somewhat arbitrary because patterns are often obscured by intergradations of the principal types of vegetation and by complexities of secondary succession. Secondary vegetation and agricultural land also are present. In figure 2.7, secondary vegetation is shown as a single color, regardless of the original type of vegetation; it includes natural second-growth vegetation and induced grasslands that are used for grazing. Under agricultural land, we pooled areas used for all types of crops. Urban areas, as well as bodies of water, are shown. Mangrove (fig. 2.8). Mangroves (Rhizophora) occur where rivers or lagoons join the coast. Height varies from 3 to 8 m. They often occur in narrow stands, but these may be several kilometers long. A small, well-preserved area of mangroves is present in southern Colima. Halophytic vegetation (fig. 2.9). Halophytic vegetation often grows where concentration of salt in soils is greater than 0.5% (Schimper 1903; Stocker 1928). In Colima this type of vegetation is mainly along the border of the Cuyutlán Lagoon. The lagoon receives saline water from the Pacific Ocean and fresh water from the Armería River (Silva-Bátiz et al. 2009).

Figure 2.8. Mangrove in dry season.

Figure 2.9. Halophytic vegetation.



STUDY AREA

15

Figure 2.10. Left: Tropical deciduous forest in dry season. Right: In wet season.

Figure 2.11. Tropical semideciduous forest.

16

Tropical deciduous forest (fig. 2.10). Adjacent to areas with arid thorn-scrub vegetation, on most of the hillsides and in many of the interior valleys, there is a relatively tall (9–15 m) homogeneous forest of mostly legumes that also includes representatives of many other tropical families (Schaldach 1963). The forest is dense in many areas, with a thick undergrowth of thorny vines. Characteristic genera include Acacia (acacia), Bombax (bombax), Brosimum (breadnut tree), Calliandra (powder puff), Cassia (cassias), Ceiba (ceiba), Enterolobium (legume), Ficus (wild figs), Hura (hura wood), Lysiloma (legume), Mimosa (mimosa), and Psidium (guava). This forest is deciduous, flowering in the dry season when more than 80% of the trees are bare of leaves and sprouting abundant new greenery with the first rains. Pachycereus (columnar cactuses) occur within this type of forest (Schaldach 1963). Tropical semideciduous forest (fig. 2.11). This type of vegetation occurs at elevations of 400– 1,200 m; thus, it usually is at higher elevation than the tropical deciduous forest. This type has been highly fragmented, resulting from production of agricultural crops and livestock. In the past, the climax stage of this forest included species of very tall tropical trees (12 to >30 m) that were 30–60 cm in diameter. At present, few trees exist that are at the higher end of this size scale. Among the characteristic species of tropical semideciduous forest are breadnut tree (Brosimum alicastrum), bully (Sideroxylon cartilagineum), a cecropia (Coussapoa purpusii), hura wood (Hura polyandra), alligator wood (Guarea glabra), angelica tree (Dendropanax arboreus), palo María (Calophyllum brasiliense var. rekoi), and occasionally, a cottonwood (Populus guzmanantlensis).

BATS OF COLIMA, MEXICO

Figure 2.12. Oak forest.

The tropical semideciduous forest is homogeneous, and merges with the tropical deciduous forest on its lower margins. At the upper margins of this forest, it is abruptly replaced, in many areas, by riparian gallery forests (Schaldach 1963; Iltis and Cochrane 1995). Riparian gallery forests occupy relatively few hectares in Colima, so it was not part of the classification system we employed; some areas of gallery forest were likely subsumed within secondary-vegetation category given the level of disturbance. Oak forest (fig. 2.12). The transition to oak forest, of which there are two types, is usually rather sharp. This type of deciduous woodland, which is characterized by oaks (Quercus castanea, Q. magnoliifolia, and Q. resinosa), and acacia (Acacia pennatula), is characteristic of drier, lower elevations of 400–1,500 m in La Media Luna (located in the municipality of Coquimatlán in central Colima) and a thin fringe around the flanks of the Volcán de Fuego. There is also a semideciduous forest of taller trees at relatively moist sites (>1,500 m); characteristic trees are white oaks (Q. cassipes) and other oaks, including Q. candicans, Q. laurina, and Q. obtusata. At its lower margins, oak forest rarely exceeds 6–8 m in height and often is dense, with a thick understory of oak seedlings and saplings in some areas. On the highest ridges and in some of the high, bowl-like valleys, the oak forest becomes more open, with taller trees. Pine-oak forest (fig. 2.13). At about 1,400 m and extending upward to 2,500 m in areas around the volcanoes—Campo Uno, Cerro Grande, and El Terrero—the oak forest merges into extensive pine-oak forest comprised of mixtures of Douglas pine (Pinus douglasiana), ocote (P. maximinoi), Figure 2.13. Pine-oak forest.



STUDY AREA

17

Nicaraguan pitch pine (P. oocarpa), Mexican red oak (Quercus castanea), and other species of oaks (Q. elliptica, Q. magnoliifolia, and Q. resinosa). Trees generally are medium-sized (≤15 m in height), but very large trees do occur, such as Q. salicifolia. This type of forest is a preclimax to oak and pine climaxes. The oaks are neither moss-covered nor epiphyte-hung in this type of forest (Schaldach 1963; Iltis and Cochrane 1995). Various species of tall grass (Andropogon) occur regularly in association with the oaks and, in some areas, cover large valleys and openings. Many forbs, including mints, grow in the pine-oak forest. Secondary vegetation (fig. 2.14). The most frequent form of secondary vegetation in Colima is arid thorn scrub, which covers the coastal plain and basal slopes of coastal mountains, and reappears again on the Plains of Colima at elevations of 800–1,200 m. This type of secondary vegetation is low, semi-open scrub composed of many kinds of spiny shrubs and legumes, including amole (Ziziphus mexicana), tormentosa (Tabernaemontana tomentosa var. palmeri), prickly-ash (Zanthoxylum), cubata (Acacia cochliacantha), cudjoewood (Bonellia macrocarpa), and the legume Sphinga acatlensis. In general, the vegetation is low in height (2–3 m), and plants are in scattered groups; pitahaya cactus (Cephalocereus mezcalensis) is also characteristic of secondary vegetation (Schaldach 1963). We have included various types of secondary growth in this category, whether or not the land has been under cultivation. Agricultural land (fig. 2.15). The nature and types of native vegetation are often masked and distorted as a result of forestry and agricultural activities. Colima has experienced a rapid rate of deforestation. From 1981 to 1992, forest coverage was reduced by 25% (Flores and Gerez 1994) and, according to Palacio-Prieto et al. (2000), 42.6% of the state is dedicated to agriculture and urban areas. The latter percentage is similar to that in INEGI (2014), which indicated that 44.1% of the area in Colima is devoted to agriculture. Olson and Dinerstein (1998) noted that the ecoregion within which Colima is located is critically threatened in terms of conservation. Due to prevalent pre-Columbian methods of agriculture, many large tracts were cut over and burned each year to create new farming fields. These fields were abandoned, usually in the second or third year, and plant successions began rapidly; occasionally, it is possible to find large, rectangular patches of thorn scrub within undisturbed tropical semideciduous forest on a hillside. This is

Figure 2.14. Secondary vegetation.

18

BATS OF COLIMA, MEXICO

Figure 2.15. Agricultural lands.

mute evidence of former fields being reclaimed not by tropical semideciduous forest but by a type of secondary vegetation that thrives due to more arid conditions (Schaldach 1963). The main agricultural crops in Colima are corn (Zea mays), sugar cane (Saccharum officinarum), jícama (Pachyrhizus erosus), sweet potato (Ipomoea batatas), different kinds of chili peppers (Capsicum), and forage. Coconut palms (Cocos nucifera) cover large areas along the coast, mainly in the western and southeastern parts of the state. In several places, coconut palms are mixed with banana (Musa), mango (Mangifera indica), lime (Citrus latifolia), and orange (C. sinensis) trees. Permanent agriculture in Colima is generally associated with livestock, and in many areas, farming involves the growing of grasses, which has resulted in fragmentation. In many instances, natural vegetation remains only on borders of water bodies and along property boundaries. No type of vegetation has escaped grazing by cattle, and overgrazing regularly occurs.



Left: Coconut grove (Cocos nucifera). Right: A field of jícama (Pachyrhizus erosus) and sugar cane (Saccharum officinarum).

STUDY AREA

19

Figure 3.0. Specimen preparation in the field in 2001. Upper left: Leobardo Sánchez Vázquez. Upper right: Luz Alcántara. Lower left: Brian Amman. Lower right: Mónica Salas-Rojas prepares specimen tags and works with the field catalog.

3

METHODS

The information provided in this book comes from three main areas: our fieldwork in Colima; a survey of specimens in mammal collections in Mexico and the United States; and our search for relevant literature.

Field Studies In addition to earlier fieldwork in Colima in the 1970s and 1980s by some of us, we conducted intensive field studies in Colima from 30 December through 14 January each year from 1996–97 through 2009–2010. Four of us—Cornelio Sánchez-Hernández, María de Lourdes Romero-Almaraz, Gary Schnell, and Michael Kennedy—participated in all fourteen trips, Troy Best in eleven trips, Sara González-Pérez in seven trips, and Robert Owen in six trips. The five coleaders of the 1999–2000 trip are shown in figure 3.1, while the other two co-authors are pictured in figure 3.2. The primary goals of fieldwork in Colima were to inventory the mammalian fauna and to obtain information on the biology and natural history of the species encountered. We used a variety of methods to inventory species of bats present in Colima. Figure 3.1. Coleaders of the 1999–2000 expedition. Left to right: Mike Kennedy, María de Lourdes (Lulú) RomeroAlmaraz, Gary Schnell, Troy Best, and Cornelio SánchezHernández in the requisite photograph with the Volcán de Fuego as a backdrop.

Figure 3.2. Two of the coauthors on trips to Colima. Left: Robert Owen in 2001. Right: Sara González-Pérez in 2007.

21

Mist nets. We sampled with mist nets 2.6 m in height and 6, 9, and 12 m in length. Nets were supported at ground level (0–3 m) and placed in vegetation along edges of roads or gaps, and along edges or above bodies of water. Efforts varied over the years, mainly due to differences in numbers of participants, habitat conditions, accessibility of study sites, and the nature of studies of other mammals being conducted. From 1997 to 2003, 30–40 mist nets were used and, from 2004 to 2010, 10–15 were employed. Typically, mist nets were opened at about 1830 hours (hereafter abbreviated as h), shortly before dusk, and then checked every 10–15 min. We routinely kept nets open until 2300–2400 h, although in some situations nets remained open all night and were checked again at about 0630 h the next morning. We chose locations that allowed us to sample as many microhabitats as possible. Bats were removed from nets and placed in cloth bags closed by drawstrings. Those not collected as voucher specimens were identified, and sex and reproductive condition were recorded before release. Roosts. We sought natural roosting sites of bats—such as caves, large crevices, and mines—by asking local people and by searching habitats. We also investigated roosts in houses, storage sheds, and culverts beneath roads. We sometimes used short-handled insect nets to capture roosting bats. Collection and preservation of specimens. We preserved voucher specimens using standard procedures described by Hall (1981) and Romero-Almaraz et al. (2007). Usually we limited collections to about 20 individuals (10 males, 10 females) of abundant species from each locality and one or two specimens for rare or uncommon species. For most bats collected, we gathered information on life history and reproductive condition, preserving skins, skeletons, and tissues; we obtained ectoparasites and brain tissues from some specimens. Of 3,394 bats collected during our fourteen seasons in the field, specimens were deposited in the following museum collections: 1,313 in the Colección Nacional de Mamíferos, Universidad Nacional Autónoma de México (CNMA); 338 in the mammal collection at the Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Morelos (UAEM); 25 at the Universidad Autónoma Metropolitana, Unidad Iztapalapa (UAM-I); and 1,718 in the Mammal Collection of the Sam Noble Oklahoma Museum of Natural History, University of Oklahoma (OMNH).

Identification and Location of Specimens To confirm identifications of potentially problematic specimens, we compared our voucher material to museum specimens and also consulted descriptions of species in the literature. In species accounts, for individual species and localities, we have identified the museum or museums where relevant specimens were deposited. Museums in which voucher and other specimens we examined are preserved include the following: AMNH, American Museum of Natural History (New York, New York); BMUNLV, University of Nevada, Las Vegas (Las Vegas); CNMA, Colección Nacional de Mamíferos, Instituto de Biología, Universidad Nacional Autónoma de México (Mexico, D.F.); FMNH, Mammal Collection, Field Museum of Natural History (Chicago, Illinois); IPN, Colección de Mamíferos, Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional (Mexico, D.F.); KU, University of Kansas Natural History Museum (Lawrence); LACM, Natural History Museum of Los Angeles County (Los Angeles, California); LSUMZ, Louisiana State University Museum of Natural Science (Baton Rouge); NMNH, United States National Museum of Natural History (Washington, D.C.); OMNH, Mammal Collection, Sam Noble Oklahoma Museum of Natural History, University of Oklahoma (Norman); UAEM, Colección de Mamíferos, Centro de Investigaciones Biológicas, Universidad Autónoma del Estado de Morelos (Cuernavaca); UAM-I, Universidad Autónoma Metropolitana, Unidad Iztapalapa (Mexico, D.F.); and UM, Mammal Collection, University of Memphis (Memphis, Tennessee). In addition, we included specimens compiled in the Mammal Networked Information System (MaNIS 2012) for the following collections: LACM, Natural History Museum of Los Angeles County (Los Angeles, California; for fluid-preserved specimens); MSB, Museum of Southwestern 22

BATS OF COLIMA, MEXICO

Biology, University of New Mexico (Albuquerque); MVZ, Museum of Vertebrate Zoology, University of California at Berkeley (Berkeley); ROM, Royal Ontario Museum (Toronto, Ontario, Canada); TTU, Museum of Texas Tech University (Lubbock); UMMZ, University of Michigan Museum of Zoology (Ann Arbor); UAZ, Mammal Collection, University of Arizona (Tucson); and WNMU, Mammal Collection, Western New Mexico University (Silver City).

Measurements Measurements of bats collected in Colima are provided in appendix B for 65 of 66 species (no specimen from Colima was located for the Brazilian brown bat, Eptesicus brasiliensis). Morphometric summaries include only measurements of adults. Linear measurements of external and cranio-dental dimensions are reported in millimeters (mm) and mass in grams (g). When samples were adequate, we assessed sexual dimorphism using an analysis of variance (ANOVA) for each character separately. The following 16 measurements (mostly from Hall 1981) are reported for many of the specimens: (1) Total length. From tip of snout to tip of last caudal vertebra. (2) Tail length. From point of dorsal flexure of tail with sacrum to tip of last caudal vertebra. (3) Foot length. From posterior edge of base of calcar to tip of claw of longest toe. (4) Ear length. From notch to fleshy tip of pinna. (5) Tibia length. From proximal to distal end of tibia. (6) Mass. Amount of matter (in grams). (7) Forearm length. From elbow (tip of olecranon process) to wrist (including carpals). (8) Third-metacarpal length. Greatest length of third metacarpal. (9) Greatest skull length. From posteriormost point on occiput to anteriormost point on premaxillae (excluding incisors). (10) Condylocanine length. From posteriormost end on occipital condyles to anteriormost end of base of upper canines. (11) M  axillary-toothrow length. From anteriormost edge of base of canine to posteriormost edge of base of M3. (12) Interorbital constriction. Least distance across top of skull between the orbits (eye sockets). (13) Zygomatic breadth. Greatest breadth across zygomatic arches. (14) Braincase breadth. Greatest breadth of globular part of braincase. (15) Mandible length. Greatest length of mandible, excluding teeth. (16) Mandibular-toothrow length. From anterior edge of alveolus of first incisor or first cheek tooth (premolar or molar) to posterior edge of alveolus of last tooth. For museum specimens, the first 4 measurements (total length, tail length, foot length, and ear length), as well as mass, were transcribed from specimen tags. The other measurements were taken directly from the specimens.



METHODS

23

Accounts Accounts for families and then species are arranged systematically within family (and within subfamily for Phyllostomidae and Vespertilionidae), and then sequentially as presented in Simmons (2005). With a few exceptions due to the inclusion by Simmons of tribes, which we do not mention, this results in an alphabetical arrangement by genus and species within a family or subfamily. Scientific names, English names, and taxonomic sequence follow Simmons with the following exceptions: (1) The northern yellow-shouldered bat (Sturnira parvidens) is recognized as specifically distinct from S. lilium based on Velazco and Patterson (2013), a change endorsed by Ramírez-Pulido et al. (2014). The English name is one used by several authors when referring to the previously designated subspecies. (2) Similarly, based on assessments by Velazco and Patterson (2013), we have employed the specific name Sturnira hondurensis for Mexican populations previously referred to as S. ludovici. The English name Honduran yellow-shouldered bat is similar to that used previously when referring to the subspecies S. ludovici hondurensis. (3) The intermediate fruit-eating bat (Artibeus intermedius) is recognized as a separate species from others in the genus based on our own experience and following Wilson (1991). (4) Three species—Aztec fruit-eating bat (Dermanura azteca), pygmy fruit-eating bat (D. phaeotis), and Toltec fruit-eating bat (D. tolteca)—are treated as generically distinct from Artibeus following Owen (1987) and several subsequent authors (e.g., Baker et al. 2003; Ramírez-Pulido et al. 2014); see review by Ramírez-Pulido et al. (2005) that indicates reasons for such recognition. (5) Our treatment follows Tejedor (2005) who described Natalus lanatus as a new species separate from N. mexicanus. We have applied the English name of woolly funnel-eared bat to the former. (6) For the Mexican greater funnel-eared bat, the name Natalus mexicanus instead of N. stramineus is employed following Tejedor (2006). (7) The separation of Eumops ferox from E. glaucinus (Wagner’s bonneted bat) is recognized based on McDonough et al. (2008). The English name used for E. ferox is fierce bonneted bat. (8) Carter’s black myotis (Myotis carteri) is recognized as specifically distinct from the black myotis (M. nigricans) based on LaVal (1973a) and Bogan (1978). Individual accounts of species start with the English common name and the scientific name. The latter is followed by the authority and year—that is, the name of the person who described the taxon and the year in which it was described. If the name of the species or placement has changed (e.g., moved to a different genus) subsequent to the original description, the name of the authority and year are placed in parentheses. In each account we have provided information on the type locality (i.e., the locality where the initial or type specimen was taken) and the derivation of the name. Most accounts then include sections for description, distribution, conservation status, environmental information, diet, reproduction, activity, other observations, measurements, specimens examined, localities, and records in literature. Description, distribution, and conservation status. Descriptions are based on literature and on our observations. In the distribution section, we indicate the complete geographic distribution of the species, as well as elevational range for the species as a whole and in Colima. Conservation status, overall for the species, is from the Red List of Threatened Species (2014) of the International Union for Conservation of Nature and Natural Resources (IUCN), while the status in Mexico is that designated in the Norma Oficial Mexicana-059-2010, Secretaría de Medio Ambiente y Recursos Naturales (SEMARNAT 2010). Habitat information is based on our field observations, as well as that garnered from the literature. Environmental information. For one-half (33) of the species, data were sufficiently robust to enable assessment of differences in environmental conditions for sites where a species was known to occur versus locations where it had not been recorded. We restricted analyses to species that were present in at least 10 localities and sites where at least 3 species were recorded, resulting in evaluation of 33 species at 161 localities (for details, see González-Pérez 2012). While it is clear that some species recorded at fewer than 10 localities were likely restricted in their distribution 24

BATS OF COLIMA, MEXICO

in Colima due to environmental conditions, statistical power was judged insufficient to yield a meaningful analysis. To eliminate from consideration those sites where only limited sampling occurred, we instituted the restriction of analyzing only sites where at least 3 species were recorded, thus reducing the number of false negatives (i.e., situations where a species was present but surveying was so limited that the species was not detected). Overall, we analyzed the following 21 environmental variables: (1) elevation in meters (fig. 2.2); (2) slope, in degrees (fig. 2.3); (3) annual mean temperature, °C (fig. 2.5); (4) annual precipitation in millimeters (fig. 2.4); (5) isothermality as a percentage; (6) maximum temperature of warmest month, °C; (7) mean diurnal temperature range, °C; (8) mean temperature of coldest quarter, °C; (9) mean temperature of driest quarter, °C; (10) mean temperature of warmest quarter, °C; (11) mean temperature of wettest quarter, °C; (12) minimum temperature of coldest month, °C; (13) precipitation of coldest quarter in millimeters; (14) precipitation of driest month in millimeters; (15) precipitation of driest quarter in millimeters; (16) precipitation seasonality, coefficient of variation, as a percentage; (17) precipitation of warmest quarter in millimeters; (18) precipitation of wettest month in millimeters; (19) precipitation of wettest quarter in millimeters; (20) temperature annual range, °C; and (21) temperature seasonality, °C. While most of these environmental variables are self-explanatory, three need clarification. Isothermality (variable 5) is variable 7 divided by variable 20, with the result multiplied by 100, thus providing an index of the degree to which temperature varies over a 24-h period relative to the range of temperature for the whole year. Precipitation seasonality (variable 16) is coefficient of variation of monthly precipitation values (i.e., [standard deviation×100]/mean), thereby indicating how precipitation varies from month to month during the year. Temperature seasonality (variable 21) is the standard deviation of monthly mean temperatures multiplied by 100, thereby indexing variability of temperatures among months. Slope was digitized by Miguel A. Ortega Huerta, and the other 20 environmental variables were acquired from WorldClim (Hijmans et al. 2005). Data on slope, precipitation, and temperature variables had a spatial resolution of 30 s (e.g., 0.93 × 0.93 = 0.86 km2 at equator), the geographic coordinate system WGS 1984 (latitude/longitude coordinate reference system, not projected), and the datum D WGS 1984. For each of the 33 species evaluated, a one-way analysis of variance (ANOVA) was completed for each of the 21 environmental variables. Variables reported in species accounts are those with statistically significant differences (P < 0.05) between sites where the species was recorded and where it was not. Of the 33 species, 22 had one or more of the environmental variables we evaluated for which there was a significant difference. Diet. Much of the information on diet was gleaned from the literature. In addition, some unpublished data were provided by the authors. Reproduction. Reproductive data were recorded for many but not all specimens from Colima. Males were separated into juveniles, subadults, and adults. For adults, we tabulated whether testes were abdominal (not descended), inguinal (not fully descended), or scrotal. Measurements of testes in millimeters are summarized; in most instances, length and width of testes were available but in others only length was recorded. In the section on reproduction, we have used a shorthand notation to convey the basic data on males in an efficient way. For example, we might write the following: Male, abdominal testes, 11 (January, 5, mean = 4.4×3.2 [2×1 to 5×4], July, 6, mean = 3.4×1.8 [2×1 to 5×3]). This indicates that there were 11 males with abdominal testes, 5 from January and 6 from July. The mean length and width of the testes (in millimeters) are given for each month, followed in brackets by the range of length and width combinations. Sometimes only length was available, so there would be only a single value for the mean and single values for each end of the range. When no measurement was provided, this is designated by an nm; at times, the nm is followed by a number in curly brackets, which indicates that the measurement was not available for more than one individual.

METHODS

25

Figure 3.3. Young of lesser long-nosed bat (Leptonycteris yerbabuenae) at roost site.

Female specimens also were partitioned into juveniles, subadults, and adults. We recorded if they had been reproductively active: pregnant, lactating (nipple with secretion of milk), postlactating (showing no hair around elongated nipples), or inactive (not pregnant and nipple with no secretion of milk). For pregnant females, we recorded number of embryos and measurements of those embryos (usually length and width but sometimes only length). In presenting detailed information on females, we have used an approach similar to that employed for males (see above). The mean and range values are for embryo lengths and widths (as indicated, sometimes only length). Age was judged in part on the basis of presence of cartilage between the diaphysis and epiphyses of phalanges, the presence of which indicated a lack of complete ossification. We considered individuals to be infants when they had cartilage between the diaphysis and epiphyses and were carried by the mother. Juveniles had space between the diaphysis and epiphyses, were left in the roost, and did not fly. In subadults, the separation between the diaphysis and epiphyses had almost disappeared, and the bats could fly. In adults, no cartilage was evident. For some species, the color of pelage also differed among age groups. Information on the reproductive pattern of the species is summarized, with most coming from the literature. We indicated whether the pattern was having one estrous period/breeding period (monoestry) or more than one (polyestry). Monoestry was categorized as “seasonal” or “asynchronous.” We also distinguished between bimodal polyestry and continuous polyestry. Activity and other observations. Data on activity were obtained from the literature. In addition, observations by the authors are reported for some species. The section on other observations, when included, provides additional information of interest concerning the species. 26

BATS OF COLIMA, MEXICO

Measurements, specimens examined, localities, and records in literature. In the section on measurements for each species, we refer to appendix B and report whether sexual dimorphism was discerned for any measurements taken. Under specimens examined, we indicate the total number of specimens and museums in which they are deposited. For localities, we refer to the map showing the distribution of the species and then, by locality, specify the museums in which specimens from given localities are located. Locality numbers refer to those listed in the gazetteer (appendix A). Locality designations with an asterisk (*) indicate that specimens were from Colima, but we were unable to determine precise geographic coordinates. Locality numbers with double asterisks (**) refer to sites that were indicated as being in Colima but which are, in fact, outside the state.

Gazetteer The gazetteer (appendix A) includes all of the localities in Colima for specimens and records of bats in the literature. Geographically close localities are designated with the same number combined with a letter (e.g., 1, 1a) but in distribution maps for species they were treated as being the same. All localities are shown on the map in figures 2.1 and A.0. The first item for each locality listed indicates the municipality within which the site is located. Original units of distance recorded on labels are given, with metric equivalents provided in brackets if English units were used. In a few instances, additional or corrected information has been added in brackets.



METHODS

27

Figure 4.0. Cassie Poindexter exhibiting a Peters’s ghostfaced bat (Mormoops megalophylla) recently captured in a nearby cave, 2007.

4

ANALYSIS

Specimens in Museums and Documented in Literature In the introduction, we provided a brief history of collecting activities in Colima. Figure 4.1 summarizes those activities over the decades, indicating the number of specimens collected that were deposited in museums in Mexico, the United States, and Canada. The first specimens of bats were obtained in Colima during the decade of 1890–99. However, for the next fifty years, almost no specimens of bats were collected there, with only 12 specimens reaching museums. Collecting resumed in 1950–59, followed by the decade during which the greatest number of specimens Decade

Figure 4.1. Number of

258 258

1890–99

Collected in decade Cumulative

specimens of bats from Colima collected by decade and deposited in a museum.

1900–09 1910–19 1920–29 5

1930–39

263 7

1940–49

270 957 1,277

1950–59

2,773

1960–69

4,000 1,664

1970–79

5,664 64

1980–89

5,728 1,274

1990–99

7,002 2,096

2000–09

9,098 22

2010

9,120 49

Unknown

9,169 0

2 ,000

4,000 6,000 Number of specimens

8,000

10,000

29

were obtained (1960–69). Significant collecting transpired in 1970–79 and then again in the two decades from 1990 through 2009; the accumulation of specimens during those three periods largely reflects the efforts of members of our group. In total, 9,169 specimens of bats from Colima were in museums by the end of 2010. Specimens of bats from Colima are deposited in nineteen collections in Mexico, the United States, and Canada, with more than 78% housed in the Sam Noble Oklahoma Museum of Natural History, the Natural History Museum of Los Angeles County, and Universidad Nacional Autónoma de México. Eleven museums house 99% of specimens collected (fig. 4.2), with the largest number residing in the Sam Noble Oklahoma Museum of Natural History. This is chiefly the result of collections made by members of our group, with specimens from our fourteen expeditions (1996–97 through 2009–2010) being noteworthy. The number of specimens in the museum was enhanced by the fact that the collection at the University of Memphis, which held specimens primarily from the 1970s collected through efforts led by Michael Kennedy, was transferred permanently to the Sam Noble Oklahoma Museum of Natural History. Specimens from Colima in the Natural History Museum of Los Angeles County were garnered mainly in the late 1950s and 1960s (fig. 4.1). Those in the Universidad Nacional Autónoma de México resulted primarily from collections made by Bernardo Villa R. and our work during the fourteen expeditions that began in 1996–97 and concluded in 2009–2010. Not surprisingly, the three collections with the greatest number of specimens of bats from Colima also include the largest number of species from the state (fig. 4.3). The Natural History Museum of Los Angeles County houses 53 of the 66 species, while the Universidad Nacional Autónoma de México has 52 and the Sam Noble Oklahoma Museum of Natural History 47. In addition, the University of Arizona has 35 species and the Louisiana State University Museum of Natural Science 24. Together, these five museums contain 65 of the 66 species of bats from Colima. Figure 4.2. Distribution of specimens of bats from Colima in 19 collections of mammals in Mexico, the United States, and Canada. Acronyms refer to the following (top to bottom): (OMNH) Sam Noble Oklahoma Museum of Natural History; (LACM) Natural History Museum of Los Angeles County; (CNMA) Universidad Nacional Autónoma de México; (UAZ) University of Arizona; (UAEM) Universidad Autónoma del Estado de Morelos; (NMNH) United States National Museum of Natural History; (IPN) Instituto Politécnico Nacional; (KU) University of Kansas Natural History Museum; (LSUMZ) Louisiana State University Museum of Natural Science; (ROM) Royal Ontario Museum; (AMNH) American Museum of Natural History; (WNMU) Western New Mexico University; (UAM-I) Universidad Autónoma Metropolitana, Unidad Iztapalapa; (FMNH) Field Museum of Natural History; (TTU) Museum of Texas Tech University; (MSB) Museum of Southwestern Biology; (UMMZ) University of Michigan Museum of Zoology; (MVZ) Museum of Vertebrate Zoology; (BMUNLV) University of Nevada, Las Vegas.

30

Museum OMNH LACM CNMA UAZ UAEM NMNH IPN KU LSUMZ ROM AMNH WNMU UAM-I FMNH TTU MSB UMMZ MVZ BMUNLV

BATS OF COLIMA, MEXICO

3,012 2,452 1,755 615 338 256 170 164 148 90 76 26 25 11 11 8 6 5 1 0

500

1,000

1,500 2,000 Number of specimens

2,500

3,000

3,500

Specimens of the following eight species from Colima are exclusively in the Natural History Museum of Los Angeles County, most of them of the family Vespertilionidae: velvety fruit-eating bat (Enchisthenes hartii), Argentinian brown bat (Eptesicus furinalis), hoary bat (Lasiurus cinereus), Townsend’s big-eared bat (Corynorhinus townsendii), southwestern myotis (Myotis auriculus), fringed myotis (M. thysanodes), cave myotis (M. velifer), and Yuma myotis (M. yumanensis). Three species—the woolly funnel-eared bat (Natalus lanatus), broad-eared free-tailed bat (Nyctinomops laticaudatus), and Mexican big-eared bat (Corynorhinus mexicanus)—are only in the Sam Noble Oklahoma Museum of Natural History. The pocketed free-tailed bat (Nyctinomops femorosaccus) is represented only in the Louisiana State University Museum of Natural Science and the Brazilian brown bat (Eptesicus brasiliensis) only in the Universidad Nacional Autónoma de México. All other species from Colima are deposited more broadly, in more than one museum. As indicated, documentation of species occurring in Colima through specimens has resulted in 66 species now known from the state (table 1.1). The initial collections in Colima (1890–92) included 18 species. During 1935, 1941, and 1956, 4 more species were added to the list for a total of 22. The greatest number of records of new species for Colima occurred during 1958–62, when 26 additional species were documented by personnel associated with the Universidad Nacional Autónoma de México, Natural History Museum of Los Angeles County, and University of Arizona, bringing the total to 48 species. Continued effort by researchers associated with the Natural History Museum of Los Angeles County and the University of Arizona in 1965, 1968, and 1969 resulted in documentation of 6 additional species, for a total of 54. The next significant changes occurred during 1972–77, when specimens were procured by the Universidad Nacional Autónoma de México, Natural History Museum of Los Angeles County, and Sam Noble Oklahoma Museum of Natural History, which resulted in 8 new species of bats for the state. Tellez-Giron et al. (1997) indicated they collected an additional species, the Brazilian brown bat (Eptesicus brasiliensis), in Colima and deposited 3 specimens in the Colección Nacional de Mamíferos; however, those Museum

Figure 4.3. Distribution of

53 52

LACM CNMA OMNH UAZ LSUMZ IPN NMNH UAEM KU AMNH WNMU ROM UAM-I TTU UMMZ FMNH MSB MVZ BMUNLV

species of bats from Colima in 19 collections of mammals in Mexico, the United States, and Canada. Museums associated with acronyms are identified in the caption for figure 4.2.

47 35 24 22 20 16 15 15 13 11 8 7 4 3 3 3 1 0



10

20

30 Number of species

40

50

60

ANALYSIS

31

Table 4.1. SPECIES OF BATS RECORDED IN COLIMA Species of bats recorded in Colima, showing year first specimen was collected, museum where that specimen resides, and first publication indicating occurrence of the species in Colima

TAXON

YEAR (MUSEUMa)

CITATION

Pteronotus davyi (Davy’s naked-backed bat)

1890 (NMNH)

Allen 1890

Balantiopteryx plicata (gray sac-winged bat)

1892 (NMNH)

Sanborn 1937

Saccopteryx bilineata (greater sac-winged bat)

1892 (NMNH)

Sanborn 1937

Desmodus rotundus (common vampire bat)

1892 (NMNH)

Ortega C. and Massieu H. 1963; Villa-R. and Alvarez L. 1963

Glossophaga soricina (Pallas’s long-tongued bat)

1892 (NMNH)

Miller 1913b

Leptonycteris nivalis (Mexican long-nosed bat)

1892 (FMNH)

Miller 1900

Leptonycteris yerbabuenae (lesser long-nosed bat)

1892 (FMNH, NMNH)

Hoffmeister 1957

Musonycteris harrisoni (banana bat)

1892 (NMNH)

Schaldach and McLaughlin 1960

Macrotus waterhousii (Waterhouse’s leaf-nosed bat)

1892 (NMNH)

Rehn 1904c

Micronycteris microtis (common big-eared bat)

1892 (NMNH)

Miller 1898

Carollia subrufa (gray short-tailed bat)

1892 (NMNH)

Hahn 1907

Artibeus jamaicensis (Jamaican fruit-eating bat)

1892 (NMNH)

Handley 1966

Dermanura phaeotis (pygmy fruit-eating bat)

1892 (FMNH, NMNH)

Andersen 1906

Pteronotus parnellii (common mustached bat)

1892 (NMNH)

Miller 1902

Noctilio leporinus (greater bulldog bat)

1892 (NMNH)

Kennedy et al. 1984

Molossus rufus (black mastiff bat)

1892 (NMNH)

Miller 1902

Tadarida brasiliensis (Brazilian free-tailed bat)

1892 (NMNH)

Shamel 1931

Rhogeessa parvula (little yellow bat)

1892 (NMNH)

Miller 1897

Mormoops megalophylla (Peters’s ghost-faced bat)

1935 (UMMZ)

Davis and Carter 1962

Natalus mexicanus (Mexican greater funnel-eared bat)

1935 (UMMZ)

Villa-R. 1966

Pteronotus personatus (Wagner’s mustached bat)

1941 (CNMA)

Anderson 1956

Molossus sinaloae (Sinaloan mastiff bat)

1956 (AMNH)

Villa-R. 1966

Glossophaga commissarisi (Commissaris’s long-tongued bat)

1958 (LACM)

Gardner 1962c

Artibeus intermedius (intermediate fruit-eating bat)

1958 (CNMA)

Villa-R. 1966

Artibeus lituratus (great fruit-eating bat)

1958 (LACM)

Villa-R. 1966

Dermanura tolteca (Toltec fruit-eating bat)

1958 (LACM)

Villa-R. 1966

1959 (CNMA, LACM)

Villa-R. 1966

Glossophaga leachii (Gray’s long-tongued bat)

1959 (LACM)

Webster 1983

Glossophaga morenoi (western long-tongued bat)

1959 (LACM)

Polaco et al. 1992

Glyphonycteris sylvestris (tricolored big-eared bat)

1959 (LACM, UAZ)

Villa-R. 1966

1959 (LACM)

Villa-R. 1966

1959 (CNMA, LACM, UAZ)

Peterson 1963

Chiroderma salvini (Salvin’s big-eyed bat)

1959 (LACM)

Villa-R. 1962

Lasiurus intermedius (northern yellow bat)

1959 (LACM, UAZ)

Anoura geoffroyi (Geoffroy’s tailless bat)

Sturnira hondurensis (Honduran yellow-shouldered bat) Sturnira parvidens (northern yellow-shouldered bat)

32

BATS OF COLIMA, MEXICO

Gardner 1962a

TAXON

Myotis fortidens (cinnamon myotis)

YEAR (MUSEUMa)

CITATION

1959 (LACM, UAZ)

Villa-R. 1966

1959 (LACM)

Hayward 1970

Eumops ferox (fierce bonneted bat)

1960 (UAZ)

Gardner 1962a

Nyctinomops aurispinosus (Peale’s free-tailed bat)

1960 (UAZ)

Gardner 1962b

Myotis carteri (Carter’s black myotis)

1960 (UAZ)

Gardner 1962a

Myotis velifer (cave myotis)

Choeronycteris mexicana (Mexican long-tongued bat)

1961 (AMNH, CNMA)

Villa-R. 1966

Molossus molossus (Pallas’s mastiff bat)

1961 (CNMA)

Álvarez and Aviña 1964

Eptesicus fuscus (big brown bat)

1961 (LACM)

Sánchez-Hernández et al. 2002

Lasiurus blossevillii (western red bat)

1961 (LACM)

Kennedy et al. 1984

Myotis californicus (Californian myotis)

1961 (LACM)

Bogan 1978

Centurio senex (wrinkle-faced bat)

1962 (CNMA, UAZ)

Villa-R. 1966

Eumops underwoodi (Underwood’s bonneted bat)

1962 (CNMA, UAZ)

Álvarez and Aviña 1964

Nyctinomops macrotis (big free-tailed bat)

1962 (CNMA, UAZ)

Villa-R. 1966

Promops centralis (big crested mastiff bat)

1962 (CNMA, UAZ)

Álvarez and Aviña 1964

Nyctinomops femorosaccus (pocketed free-tailed bat)

1965 (LUSMZ)

Sánchez-Hernández et al. 2002

Eptesicus furinalis (Argentinian brown bat)

1968 (LACM)

Sánchez-Hernández et al. 2002

Corynorhinus townsendii (Townsend’s big-eared bat)

1968 (LACM)

Sánchez-Hernández et al. 2002

Dermanura azteca (Aztec fruit-eating bat)

1969 (LACM, OMNH)

Kennedy et al. 1984

Cynomops mexicanus (Mexican dog-faced bat)

1969 (LACM)

Sánchez-Hernández et al. 2002

Myotis auriculus (southwestern myotis)

1969 (LACM)

Sánchez-Hernández et al. 2002

Natalus lanatus (woolly funnel-eared bat)

1972 (OMNH)

Enchisthenes hartii (velvety fruit-eating bat)

1975 (LACM)

Arroyo-Cabrales and Owen 1996

Nyctinomops laticaudatus (broad-eared free-tailed bat)

1975 (OMNH)

Best and Kennedy 1984, Kennedy et al. 1984

Lasiurus cinereus (hoary bat)

1975 (LACM)

Sánchez-Hernández et al. 2002

Myotis melanorhinus (dark-nosed small-footed myotis)

1975 (LACM)

Bogan 1978

Myotis thysanodes (fringed myotis)

1975 (LACM)

Sánchez-Hernández et al. 2002

Myotis yumanensis (Yuma myotis)

1975 (LACM)

Sánchez-Hernández et al. 2002

Diclidurus albus (northern ghost bat)

1977 (CNMA)

Sánchez Hernández and Chávez Tapia 1984

Eptesicus brasiliensis (Brazilian brown bat)

1992 (CNMA)b

Tellez-Giron et al. 1997

Corynorhinus mexicanus (Mexican big-eared bat)

1997 (OMNH)

Sánchez-Hernández et al. 2002

Myotis albescens (silver-tipped myotis)

1998 (CNMA, OMNH)

Sánchez-Hernández et al. 2002

Lasiurus xanthinus (western yellow bat)

2000 (CNMA, OMNH)

Sánchez-Hernández et al. 2002

a Museums: (AMNH) American Museum of Natural History; (CNMA) Universidad Nacional Autónoma de México; (FMNH) Field Museum of Natural History; (LACM) Natural History Museum of Los Angeles County; (LSUMZ) Louisiana State University Museum of Natural Science; (NMNH) United States National Museum of Natural History; (OMNH) Sam Noble Oklahoma Museum of Natural History; (UAZ) University of Arizona; (UMMZ) University of Michigan Museum of Zoology. b

Year specimens collected not indicated in publication but likely in early 1990s. Specimens have not been located.



ANALYSIS

33

specimens have not been located. Our efforts in 1996–2010 resulted in records of an additional 3 species of bats previously unknown from Colima, for a total of 66 species. The earliest specimen of a bat in a museum from Colima was a Davy’s naked-backed bat (Pteronotus davyi) collected on 15 January 1890 on the Plains of Colima by Audley C. Buller, which was reported in December of that year by J. A. Allen (1890). Subsequently, the list of known species from Colima based on the first museum specimens taken grew at an uneven pace over the next 110 years, with reports in the literature of new species for Colima not surprisingly lagging behind procurement of the initial voucher specimens (fig. 4.4). As shown in figure 4.4 (with details in table 4.1), the published list of species known for Colima grew in spurts, as reflected in a series of papers in 1962 by Gardner (1962a, 1962b, 1962c), the 1966 compendium on bats of Mexico by Villa-R. (1966), the list of records documented in 1984 by Kennedy et al. (1984), and the work in 2002 by Sánchez-Hernández et al. (2002). One species documented in this book, the woolly funnel-eared bat (Natalus lanatus), has not previously been reported in Colima. For the period 1890–2010, specimens were obtained during 47 of those years. The degree of effort expended to obtain specimens varied substantially from year to year, as did the methods used. The species encountered during the greatest number of years were Pallas’s long-tongued bat (Glossophaga soricina, 37 years), common vampire bat (Desmodus rotundus, 31), Jamaican fruit-eating bat (Artibeus jamaicensis, 31), common mustached bat (Pteronotus parnellii, 31), little yellow-shouldered bat (Sturnira parvidens, 29), gray sac-winged bat (Balantiopteryx plicata, 29), pygmy fruit-eating bat (Dermanura phaeotis, 28), and Toltec fruit-eating bat (D. tolteca, 28). At the other extreme, there were 7 species, each of which were collected in only one year: velvety fruit-eating bat (Enchisthenes hartii), pocketed free-tailed bat (Nyctinomops femorosaccus), Brazilian brown Figure 4.4. Cumulative number

70

of known species of bats from Colima, by year, based on museum specimens and as reported in the literature.

65

Known from specimens Recorded in literature

60 55 50

Number of species

45 40 35 30 25 20 15 10 5 0 1890

34

BATS OF COLIMA, MEXICO

1900

1910

1920

1930

1940 1950 Year

1960

1970

1980

1990

2000

Family Emballonuridae

Figure 4.5. (A) Number of species per family. (B) Number of museum specimens per family.

A

3

Phyllostomidae

25

Mormoopidae

4

Noctilionidae

1

Natalidae

2

Molossidae

12

Vespertilionidae

19 0

5

10

15 Number of species

20

25

30

Family Emballonuridae

B

598

Phyllostomidae

6,604

Mormoopidae

1,034

Noctilionidae

18

Natalidae

150

Molossidae

437

Vespertilionidae

328 0

1 ,000

2,000

3,000 4,000 Number of specimens

5,000

6,000

7,000

bat (Eptesicus brasiliensis), hoary bat (Lasiurus cinereus), Mexican big-eared bat (Corynorhinus mexicanus), fringed myotis (Myotis thysanodes), and Yuma myotis (M. yumanensis). Within the 7 families of bats in Colima, the number of species per family varies from 1 in Noctilionidae to 25 in Phyllostomidae (fig. 4.5A). Over 70% of specimens in museums represent the family Phyllostomidae (fig. 4.5B). The Jamaican fruit-eating bat is the species most frequently captured and preserved, with fully 1,654 specimens (18% of total specimens of bats for Colima) in museums.

Natural History Diet. Bats in Colima are diverse in what they eat (fig. 4.6, table 4.2). While more than 60% (42) of species feed primarily on insects, there are 9 species that show preference for pollen, while sometimes eating other plant or animal material. Among 13 species that are frugivores, 8 also eat other types of food, while diets of the other 5 frugivore species are largely limited to fruits. The species within the bat fauna for Colima with unusual diets are the single species that is a piscivore (greater bulldog bat, Noctilio leporinus) and a sanguivore, feeding primarily on blood (common vampire bat, Desmodus rotundus). The family of bats with the greatest diversity of diets is Phyllostomidae,



ANALYSIS

35

Figure 4.6. Diets of bats in Colima.

Insectivore

42

Pollenivore-Insectivore

9

Frugivore-Omnivore

8

Frugivore

5

Piscivore

1

Sanguivore

1 0

10

20

30

40

50

Number of species which includes all types of diets except piscivory. All species in Colima of families Emballonuridae, Mormoopidae, Natalidae, Vespertilionidae, and Molossidae are insectivores. Reproductive pattern. Almost 60% of the species of bats in Colima are seasonal monoestrous (fig. 4.7, table 4.3), with mating typically taking place from January through March, and parturition from May through July. Weaning occurs at 3–9 weeks of age. These species feed on insects, which are more abundant during the rainy season (June–September), and young are weaned during the time of greatest abundance of food (July–August). Of the species, more than 25% exhibit bimodal polyestry (fig. 4.7), having two periods of mating and birthing each year; parturition usually occurs at the beginning of the calendar year and then again at the middle or near the end of the year. Species with this reproductive pattern consume fish, fruit, nectar, pollen, or insects, and some exhibit seasonal migration in search of food and roosts. The reproductive pattern of 9 species is continuous polyestry, with 3 birthing periods per year. These include 2 species of Glossophaga (Rasweiler 1972, 1974; Wilson 1979; Sánchez Hernández and Romero Almaraz 1995a), 1 species of Sturnira (Sánchez Hernández et al. 1986), all 3 species of Artibeus (Sánchez Hernández et al. 1990a; Sánchez Hernández and Romero Almaraz 1995a), 2 species of Dermanura (Sánchez Hernández and Romero Almaraz 1995a), and the 1 species of Molossus (Alberico et al. 2005). In dietary habits, species with continuous polyestry are mostly frugivore-omnivore (5 species), but 2 are pollenivore-insectivores and 1 is a frugivore (tables 4.2 and 4.3). Because production of fruits and flowers occurs at various times throughout the year in tropical regions, these bats do not often encounter significant limitations in availability of food, although several species exhibit local movements in search of food. Their young may be born during any season. Young of these species are generally precocial, with a growth period of three weeks to two months. The reproductive pattern of the common vampire bat (Desmodus rotundus) is asynchronous monoestry. The species mates and gives birth at any time during the year. Figure 4.7. Reproductive patterns of bats in Colima.

39

Seasonal monoestry Bimodal polyestry

17

Continuous polyestry

9

Asynchronous monoestry

1 0

10

20

Number of species 36

BATS OF COLIMA, MEXICO

30

40

Table 4.2. DIETS OF BATS IN COLIMA INSECTIVORE (n = 42; 63.6%) Balantiopteryx plicata (gray sac-winged bat) Diclidurus albus (northern ghost bat) Saccopteryx bilineata (greater sac-winged bat) Glyphonycteris sylvestris (tricolored big-eared bat) Macrotus waterhousii (Waterhouse’s leaf-nosed bat) Mormoops megalophylla (Peters’s ghost-faced bat) Pteronotus davyi (Davy’s naked-backed bat) Pteronotus parnellii (common mustached bat) Pteronotus personatus (Wagner’s mustached bat) Natalus lanatus (woolly funnel-eared bat) Natalus mexicanus (Mexican greater funnel-eared bat) Cynomops mexicanus (Mexican dog-faced bat) Eumops ferox (fierce bonneted bat) Eumops underwoodi (Underwood’s bonneted bat) Molossus molossus (Pallas’s mastiff bat) Molossus rufus (black mastiff bat) Molossus sinaloae (Sinaloan mastiff bat) Nyctinomops aurispinosus (Peale’s free-tailed bat) Nyctinomops femorosaccus (pocketed free-tailed bat) Nyctinomops laticaudatus (broad-eared free-tailed bat) Nyctinomops macrotis (big free-tailed bat) Promops centralis (big crested mastiff bat) Tadarida brasiliensis (Brazilian free-tailed bat) Eptesicus brasiliensis (Brazilian brown bat) Eptesicus furinalis (Argentinian brown bat) Eptesicus fuscus (big brown bat) Lasiurus blossevillii (western red bat) Lasiurus cinereus (hoary bat) Lasiurus intermedius (northern yellow bat) Lasiurus xanthinus (western yellow bat) Rhogeessa parvula (little yellow bat) Corynorhinus mexicanus (Mexican big-eared bat) Corynorhinus townsendii (Townsend’s big-eared bat) Myotis albescens (silver-tipped myotis) Myotis auriculus (southwestern myotis) Myotis californicus (Californian myotis) Myotis carteri (Carter’s black myotis) Myotis fortidens (cinnamon myotis) Myotis melanorhinus (dark-nosed small-footed myotis) Myotis thysanodes (fringed myotis) Myotis velifer (cave myotis) Myotis yumanensis (Yuma myotis)



POLLENIVORE-INSECTIVORE (n = 9; 13.6%) Anoura geoffroyi (Geoffroy’s tailless bat) Choeronycteris mexicana (Mexican long-tongued bat) Glossophaga commissarisi (Commissaris’s long-tongued bat) Glossophaga leachii (Gray’s long-tongued bat) Glossophaga morenoi (western long-tongued bat) Glossophaga soricina (Pallas’s long-tongued bat) Leptonycteris nivalis (Mexican long-nosed bat) Leptonycteris yerbabuenae (lesser long-nosed bat) Musonycteris harrisoni (banana bat) FRUGIVORE-OMNIVORE (n = 8; 12.1%) Micronycteris microtis (common big-eared bat) Carollia subrufa (gray short-tailed bat) Sturnira hondurensis (Honduran yellow-shouldered bat) Sturnira parvidens (northern yellow-shouldered bat) Artibeus intermedius (intermediate fruit-eating bat) Artibeus jamaicensis (Jamaican fruit-eating bat) Artibeus lituratus (great fruit-eating bat) Dermanura phaeotis (pygmy fruit-eating bat) FRUGIVORE (n = 5; 7.6%) Centurio senex (wrinkle-faced bat) Chiroderma salvini (Salvin’s big-eyed bat) Dermanura azteca (Aztec fruit-eating bat) Dermanura tolteca (Toltec fruit-eating bat) Enchisthenes hartii (velvety fruit-eating bat) PISCIVORE (n = 1; 1.5%) Noctilio leporinus (greater bulldog bat) SANGUIVORE (n = 1; 1.5%) Desmodus rotundus (common vampire bat)

ANALYSIS

37

Zoogeographic Associations The most abundant and diverse family of bats centered in the Neotropics is Phyllostomidae, which is represented in Colima by 25 species (table 1.1). The most diverse Nearctic family is Vespertilionidae, which is represented by 19 species in Colima. The relatively high proportions of both of these families demonstrate the transitional character of the state of Colima, between the Nearctic and Neotropical biogeographic regions of the Americas. In terms of higher taxonomic composition, bats in Colima represent a typical Neotropical lowland fauna. All seven families common in Mexico are represented. The one family in Mexico not documented for Colima is Thyropteridae (disc-winged bats), a small family known in Mexico only from Veracruz and Chiapas (Villa-R. 1966).

Conservation and Endemic Species In the Norma Oficial Mexicana-059–2010 (SEMARNAT 2010), 8 (12.1%) of the 66 species in Colima are listed as being of special concern. One species, the banana bat (Musonycteris harrisoni), is characterized as endangered. Three considered to be threatened are the Mexican long-tongued bat (Choeronycteris mexicana), Mexican long-nosed bat (Leptonycteris nivalis), and lesser long-nosed bat (L. yerbabuenae). Four species—the velvety fruit-eating bat (Enchisthenes hartii), Mexican dog-faced bat (Cynomops mexicanus), silver-tipped myotis (Myotis albescens), and Carter’s black myotis (Myotis carteri [as M. nigricans])—have been afforded protected status. On the IUCN Red List of Threatened Species (IUCN 2014), the Mexican long-nosed bat is designated as endangered, and the lesser long-nosed bat and banana bat are judged to be vulnerable. Two species are considered to be near-threatened: the Mexican long-tongued bat and Mexican bigeared bat (Corynorhinus mexicanus). The other 61 species of bats in Colima have been designated as being of least concern. Of the 66 species of bats in Colima, 5 (7.6%) are endemic to Mexico (Simmons 2005). Three of these (as indicated above) are considered to be of special concern (SEMARNAT 2010; IUCN 2014): banana bat, Mexican big-eared bat, and Carter’s black myotis. The other endemic species are the western long-tongued bat (Glossophaga morenoi) and the little yellow bat (Rhogeessa parvula).

Misidentified Species from Colima When Andersen (1906) described the hairy fruit-eating bat (Artibeus hirsutus), he cited three specimens from Colima deposited in the United States National Museum of Natural History. Specimens of this species from Colima are not in the catalog of the museum, but there is a skull of a Jamaican fruit-eating bat (A. jamaicensis) with the name of Artibeus hirsutus crossed out with pencil. It is likely that this skull was among the three skulls cited by Andersen (1906) as A. hirsutus and later changed to A. jamaicensis. During the course of examining species of bats from Colima, we scrutinized 2,402 specimens of Artibeus, none of which was A. hirsutus. In addition to recording two specimens from Colima of Carollia castanea subrufa (now C. subrufa, gray short-tailed bat, for populations along the Pacific coast of Mexico), Villa-R. (1966) identified eight specimens as Seba’s short-tailed bat (Carollia perspicillata) from a locality in the state (El Mixcoate, 9 km W Pueblo Juárez, 1,347 m). In our database, we have ten specimens of C. subrufa collected in 1961 by B. Villa-R. from that same locality. In Villa-R. (1966:268), this locality on the distribution map for C. perspicillata is notably north of other specimen localities he recorded for this species along or close to the Pacific coast. Subsequent authors (e.g., Pine 1972; Cloutier and Thomas 1992; Simmons 2005) have judged the species not to be present along the Pacific coast north of Oaxaca. We consider the purported record of C. perspicillata for Colima to be in error, with the specimens in question being C. subrufa.

38

BATS OF COLIMA, MEXICO

Table 4.3. REPRODUCTIVE PATTERNS OF BATS IN COLIMA SEASONAL MONOESTRY (n = 39; 59.1%)

BIMODAL POLYESTRY (n = 17; 25.8%)

Balantiopteryx plicata (gray sac-winged bat)

Glossophaga commissarisi (Commissaris’s long-tongued bat)

Diclidurus albus (northern ghost bat)

Glossophaga morenoi (western long-tongued bat)

Saccopteryx bilineata (greater sac-winged bat)

Leptonycteris yerbabuenae (lesser long-nosed bat)

Anoura geoffroyi (Geoffroy’s tailless bat)

Musonycteris harrisoni (banana bat)

Choeronycteris mexicana (Mexican long-tongued bat)

Macrotus waterhousii (Waterhouse’s leaf-nosed bat)

Leptonycteris nivalis (Mexican long-nosed bat)

Carollia subrufa (gray short-tailed bat)

Glyphonycteris sylvestris (tricolored big-eared bat)

Sturnira hondurensis (Honduran yellow-shouldered bat)

Micronycteris microtis (common big-eared bat)

Centurio senex (wrinkle-faced bat)

Mormoops megalophylla (Peters’s ghost-faced bat)

Chiroderma salvini (Salvin’s big-eyed bat)

Pteronotus davyi (Davy’s naked-backed bat)

Dermanura azteca (Aztec fruit-eating bat)

Pteronotus parnellii (common mustached bat)

Enchisthenes hartii (velvety fruit-eating bat)

Pteronotus personatus (Wagner’s mustached bat)

Noctilio leporinus (greater bulldog bat)

Natalus lanatus (woolly funnel-eared bat)

Molossus rufus (black mastiff bat)

Natalus mexicanus (Mexican greater funnel-eared bat)

Molossus sinaloae (Sinaloan mastiff bat)

Cynomops mexicanus (Mexican dog-faced bat)

Eptesicus furinalis (Argentinian brown bat)

Eumops ferox (fierce bonneted bat)

Rhogeessa parvula (little yellow bat)

Eumops underwoodi (Underwood’s bonneted bat)

Myotis albescens (silver-tipped myotis)

Nyctinomops aurispinosus (Peale’s free-tailed bat) Nyctinomops femorosaccus (pocketed free-tailed bat)

CONTINUOUS POLYESTRY (n =9; 13.6%)

Nyctinomops laticaudatus (broad-eared free-tailed bat)

Glossophaga leachii (Gray’s long-tongued bat)

Nyctinomops macrotis (big free-tailed bat)

Glossophaga soricina (Pallas’s long-tongued bat)

Promops centralis (big crested mastiff bat)

Sturnira parvidens (northern yellow-shouldered bat)

Tadarida brasiliensis (Brazilian free-tailed bat)

Artibeus intermedius (intermediate fruit-eating bat)

Eptesicus brasiliensis (Brazilian brown bat)

Artibeus jamaicensis (Jamaican fruit-eating bat)

Eptesicus fuscus (big brown bat)

Artibeus lituratus (great fruit-eating bat)

Lasiurus blossevillii (western red bat)

Dermanura phaeotis (pygmy fruit-eating bat)

Lasiurus cinereus (hoary bat)

Dermanura tolteca (Toltec fruit-eating bat)

Lasiurus intermedius (northern yellow bat)

Molossus molossus (Pallas’s mastiff bat)

Lasiurus xanthinus (western yellow bat) Corynorhinus mexicanus (Mexican big-eared bat) Corynorhinus townsendii (Townsend’s big-eared bat)

ASYNCHRONOUS MONOESTRY (n = 1; 1.5%) Desmodus rotundus (common vampire bat)

Myotis auriculus (southwestern myotis) Myotis californicus (Californian myotis) Myotis carteri (Carter’s black myotis) Myotis fortidens (cinnamon myotis) Myotis melanorhinus (dark-nosed small-footed myotis) Myotis thysanodes (fringed myotis) Myotis velifer (cave myotis) Myotis yumanensis (Yuma myotis)



ANALYSIS

39

Figure 5.0. Cornelio SánchezHernández working to extricate an intermediate fruit-eating bat (Artibeus intermedius) from a mist net in which it became entangled, 2006.

5

KEY TO BATS OF COLIMA

This identification key is for the 66 species that are known to occur in Colima (table 1.1). The first part of the key distinguishes among families of those bats. The family Phyllostomidae is further separated into subfamilies. Within families (and subfamilies of Phyllostomidae), the key proceeds to distinguish genera and then species. Characteristics used in the key include external, dental, and skull characteristics; in numerous cases, only a subset of the characteristics mentioned is needed for a positive identification. The following general diagrams are helpful when using the key: figure 5.1 shows general external anatomy of a bat; figure 5.2 indicates skull measurements; figure 5.3 depicts the dentition; and figure 5.4 identifies bones of the skull of a young specimen. Rostrum width (the maximal width of the rostrum), not defined elsewhere, is also used in the key. When species identification is determined within the key, reference is given to the photograph of the species; in addition, skull photographs that depict particular characteristics of interest are also sometimes referenced in the key. Note that one species, the silver-tipped myotis (Myotis albescens), appears twice in the key due to intraspecific variability.

Figure 5.1. External anatomy of a bat.

Metacarpal 3rd digit 1st phalanx 2nd phalanx 3rd phalanx

1st digit (thumb)

Tragus Noseleaf Ear

Metacarpal (2nd digit) Forearm

Metacarpal 4th digit 1st phalanx 2nd phalanx 3rd phalanx Metacarpal 1st phalanx 5th digit 2nd phalanx 3rd phalanx

Propatagium Dactylopatagium Plagiopatagium

Femur Tibia Tail

Uropatagium Calcar

41

Figure 5.2. (A) Dorsal, (B) ventral, and (C) lateral views of cranium plus (D) lateral view of mandible. Abbreviations: (BCB) braincase breadth, (CCL) condylocanine length, (GSL) greatest skull length, (IC) interorbital constriction, (MST) mastoid breadth, (MDL) mandible length, (MDTR) mandibulartoothrow length, (MTR) maxillary-toothrow length, (RB) rostral breadth, (ZB) zygomatic breadth. Redrawn from Sánchez Hernández and Romero Almaraz (1995a).

A

BCB

IC

B

MST

RB

C

MTR

D

CCL GSL

MDTR

MDL Figure 5.3. Dentition of generalized bat showing placement of individual teeth in (A) left maxilla and (B) left mandible. Abbreviations for individual teeth as follows: (I3) upper incisor, (C) upper canine, (P3, P4) upper premolars, (M1, M2, M3) upper molars, (i1, i2) lower incisors, (c) lower canine, (p3, p4) lower premolars, and (m1, m2, m3) lower molars. Numbers refer to basic structures on cheek teeth: (1) cingulum, (2) parastyle, (3) mesostyle, (4) metastyle, (5) procone, (6) hypocone, (7) paracone, (8) metacone, (9) paraconid, (10) protoconid, (11) metaconid, (12) hypoconid, and (13) entoconid. In this example, dental formula is i 1/2, c 1/1, p 2/2, m 3/3, total 30. Redrawn from Hall (1981).

42

BATS OF COLIMA, MEXICO

A

P4

2 3 M1 4

M2

M3

P3

C I3 1

B

i1

5 11 12

i2 c

p3

p4

7

6

8

13

9 m1 10

m2

m3

ZB

Figure 5.4. Basic components of skull of young bat in (A) dorsal, (B) ventral, and (C) lateral views of cranium plus (D) lateral view of mandible. Lowercase letters indicate location of bones and structures: (a) premaxilla, (b) nasal, (c) maxilla, (d) frontal, (e) parietal, (f), occipital, (g) zygomatic arch, (h) incisive foramen, (i) palatine, (j) presphenoid, (k) basisphenoid, (l) basioccipital, (m) foramen magnum, (n) auditory bulla, (o) temporal, (p) dentary, (q) coronoid process, (r) condyloid process, and (s) angular process.

A a b

e

d

c

f

g

g

B

a

c

h

j

i

k l

m

f

n

C

d e b a

o c

f

g n q

D

r p

s

Families 1  a) Noseleaf present, having spear (fig. 5.5A) or horseshoe shape (in Desmodus, fig. 5.5B), and modified in Centurio (fig. 6.47); tail vertebrae present or absent (fig. 5.5C and 5.5D); dentition variable in size and form (upper molars with W-shape in subfamily Phyllostominae, fig. 5.5E); total number of teeth = 20–34 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllostomidae 9  b) Noseleaf absent; tail vertebrae present; dentition with some variability in form (upper molars W-shaped); total number of teeth = 26–38 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 a) Forearm length >80 mm; upper lip divided or “leporinus” (fig. 5.5F); feet and claws greatly enlarged and flattened laterally; mastoid region conspicuously flared and shelflike; total number of teeth = 28 . . . . Noctilionidae . . . . . Noctilio leporinus (greater bulldog bat, fig. 6.69)  b) Forearm length 7 mm beyond margin of uropatagium (fig. 5.6B); ears bonnet-shaped and often united across forehead (fig. 5.6C); premaxillae with nasal branches present; palatal branches, when present, form 2 palatal foramina and, when absent, only 1 foramen present; total number of teeth = 26–30 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molossidae 37 b) Tail well developed, extending to edge of uropatagium or few millimeters beyond (fig. 5.6D); ears variable in form and size but not bonnet-shaped; palatal branches of premaxillae, if present, are bony and, if absent, the palate is widely emarginated anteriorly; total number of teeth = 30–38 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Figure 5.6. Characteristics

A

B

C

D

E

F

G

H



of bats: (A) front of skull pointed downward, with top arrow indicating position of postorbital process behind very small orbit, and lower arrow pointing to greatly inflated rostrum; (B) dorsal view, tail extending beyond margin of uropatagium; (C) ears bonnet-shaped and united across forehead; (D) dorsal view, tail well developed, extending to edge of uropatagium; (E) gland or natalid organ in frontal region; (F) dorsal view, male with keratin capsule dorsally on uropatagium; (G) dorsal view, female with faint marks where keratin capsule exists in male; (H) dorsal view, rostrum wide with hexagonal depression anteriorly.

KEY TO BATS OF COLIMA

45

6 a) Fragile appearance; eyes diminutive; funnel-shaped ear (fig. 6.72 and 6.74); tragus small and triangular; tibia length >50% of forearm length; males with gland or natalid organ in frontal region (fig. 5.6E); skull with long narrow rostrum; braincase globate and elevated; total number of teeth = 38 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Natalidae 36 b) Some species have fragile appearance but most do not; most species with small eyes; ears variable in form and size, but not funnel-shaped; tragus prominent (fig. 6.112); tibia length 90 mm; dorsal and ventral fur white or with gray base and white tip; no wing sac dorsally on propatagium; eyes large; males with keratin capsule dorsally on uropatagium (fig. 5.6F), females with faint marks where capsule exists in males (fig. 5.6G); rostrum wide with hexagonal depression anteriorly (fig. 5.6H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diclidurus albus (northern ghost bat, fig. 6.4)  b) Total length usually 15.5 mm . . . . . . . . . . . . . . . . . . . Corynorhinus townsendii (Townsend’s big-eared bat, fig. 6.120)  b) Fur gray-brown to blackish; hair of dorsum sharply bicolored; uropatagium with 17.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eptesicus fuscus (big brown bat, fig. 6.106) 54

BATS OF COLIMA, MEXICO

 b) Forearm length 14 females and a dominant male) may include an additional subordinate male (Ortega and Arita 2002). Activity. In Michoacán, most Jamaican fruit-eating bats were captured in mist nets from 1845 to 2215 h (unpublished data). After that time, activity diminished, with the species becoming active again in early morning. Some have been recaptured up to 4.7 km from where originally marked (mean = 2.9 km; Montiel et al. 2006). Other observations. Variation at nuclear microsatellite loci was analyzed to assess genetic structure, relatedness, and paternity of Jamaican fruit-eating bats (Ortega et al. 2003). Those captured in two caves 11 km apart exhibited little genetic differentiation between caves, providing no evidence of harem structure. Dominant males and subordinate males (i.e., those with lower social rank) shared paternity in large groups, while dominant males and satellite males (i.e., those not having their own territory) shared paternity in smaller groups. Results suggest that rank of a male influences paternity. The lifespan of Jamaican fruit-eating bats is 7–10 years (Jones 1982; Harvey et al. 2011). In Colima, 34 Jamaican fruit-eating bats were evaluated for antirabic antibodies (Salas-Rojas et al. 2004). For 22 in a disturbed area (primarily agricultural), 7 (31.8%) tested positive, while for 12 in an undisturbed area (tropical semideciduous forest), 7 (58.3%) tested positive. A positive test indicated that an individual had been exposed to rabies (Lyssavirus). The greater prevalence in the undisturbed area may result

Artibeus jamaicensis

113

from more frequent interspecies interactions, possibly due to sharing shelters. Measurements. Males were significantly larger than females in zygomatic breadth and mandibular-toothrow length. Females were significantly larger than males in total length, mass, forearm length, and third-metacarpal length (appendix B). Total length, obtained from specimen tags, exhibited a marked bimodal distribution in both males and females, with modes for males at 78 and 100 mm and for females at 80 and 100 mm. The bimodal distributions are not biologically meaningful but reflect differences in how investigators measure total length in species that do not have an external tail; some investigators measure to the posterior edge of the body of the bat not including the uropatagium, while others measure to the posterior edge of the uropatagium. Specimens examined (n = 1,654). 9 AMNH; 296 CNMA; 17 IPN; 8 KU; 455 LACM; 2 LSUMZ; 5 NMNH; 753 OMNH; 9 ROM; 68 UAEM; 6 UAM-I; 25 UAZ; 1 WNMU. Localities (fig. 6.44). 1d. 1 OMNH; 3a. 5 CNMA, 7 OMNH; 3c. 13 LACM; 5a. 1 LACM; 8. 1 OMNH; 12a. 13 LACM; 12f. 1 CNMA; 16c. 53 LACM; 16d. 4 ROM; 17. 6 LACM; 19. 1

OMNH; 21. 27 CNMA, 86 OMNH, 22 UAEM; 24. 3 UAM-I; 28. 1 OMNH; 29. 1 LACM; 32a. 1 LACM; 32b. 4 CNMA; 32e. 16 LACM; 32g. 13 LACM; 32j. 30 LACM; 32k. 5 LACM; 33b. 4 UAZ; 34. 3 LACM; 37. 6 LACM; 38. 3 IPN; 40. 4 CNMA, 1 UAEM; 40a. 3 CNMA; 41. 21 CNMA, 1 OMNH, 2 UAEM; 42. 2 CNMA, 5 UAEM; 43. 5 OMNH; 44. 7 LACM; 45. 5 CNMA, 5 UAEM; 47. 2 CNMA, 2 UAEM; 48. 25 CNMA; 49. 3 CNMA, 5 OMNH; 50. 4 OMNH; 50a. 23 OMNH; 51. 7 OMNH; 52. 2 CNMA, 169 OMNH; 52a. 8 OMNH; 52b. 2 OMNH; 52c. 5 OMNH; 53. 47 OMNH; 53a. 6 OMNH; 54. 1 OMNH; 54a. 1 CNMA, 92 OMNH; 56. 3 OMNH; 57. 7 LACM; 60. 2 OMNH; 60a. 2 CNMA; 62. 22 LACM; 65. 2 OMNH; 67. 1 OMNH; 68. 1 CNMA, 9 OMNH; 69. 1 OMNH; 70. 1 CNMA, 6 OMNH; 72. 2 OMNH; 72a. 1 CNMA, 2 OMNH; 73. 4 OMNH; 74. 1 OMNH; 75. 1 CNMA, 2 OMNH; 77. 1 CNMA, 6 OMNH; 78. 1 UAEM; 81. 3 CNMA, 1 OMNH; 83. 8 LACM; 84. 21 LACM; 85. 38 LACM; 85a. 2 LACM; 86. 1 UAM-I; 88. 5 CNMA; 88a. 17 CNMA, 11 OMNH; 88b. 3 CNMA, 1 OMNH, 4 UAEM; 89. 9 CNMA, 1 OMNH, 1 UAEM; 90. 6 CNMA, 1 OMNH, 1 UAEM; 91. 15 CNMA, 1 OMNH, 9 UAEM; 92. 11 CNMA; 94. 1 UAEM; 96. 1 CNMA; 97. 7 CNMA, 22 OMNH, 4 UAEM; 97a. 5 OMNH; 97b. 2 CNMA, 2 OMNH; 97c. 4

Figure 6.44. Known distribution

c

ce

O

19°30'

ifi

Copyright © 2016, University of Oklahoma Press.

JALISCO

Pac

of Jamaican fruit-eating bat (Artibeus jamaicensis) in Colima.

an

1

MEXICO

COLIMA

19

78

19°00'

8

37

38 44 40 41 42 43 29 17 28 Cuauhtémoc 45 16 47 32 33 VILLA DE Comala 49 50 52 48 53 34 ÁLVAREZ MINATITLÁN 51 56 Villa de 57 54 122 Álvarez 149 CUAUHTÉMOC 60 123 84 67 68 Colima 83 70 72 62 154 65 COQUIMATLÁN 86 85 73 69 129 138 77 89 74 75 81 90 88 130 160 135 139Coquimatlán 94 91 92 COLIMA 161 133 MANZANILLO 142 96 131 141 145 100 104 109 97 98 167 169 103 Manzanillo 168 170 114 177 113 171 115 ARMERÍA 198 116 199 179 178 180 222 Ixtlahuacán 183 IXTLAHUACÁN 184 205 193 Armería 224 175 Tecomán 206 225 195 226 TECOMÁN Minatitlán 24

19°15'

5 21

3

Jamaican Fruit-eating Bat

COMALA

12

210

Artibeus jamaicensis

18°45'

215 216

20 Kilometers 10 Miles

MICHOACÁN

N 104°45'

114

FAMILY PHYLLOSTOMIDAE

104°30'

104°15'

104°00'

103°45'

103°30'

CNMA; 97d. 2 OMNH; 97e. 3 KU; 98g. 5 OMNH; 100. 6 OMNH; 103. 7 CNMA, 1 OMNH; 104. 2 UAM-I; 109. 5 KU; 113. 12 CNMA, 2 OMNH; 114f. 1 CNMA; 114g. 3 CNMA; 114i. 7 UAZ; 115. 1 CNMA; 116. 18 OMNH; 116a. 5 CNMA, 1 OMNH; 116b. 8 CNMA, 8 OMNH; 122a. 9 LACM; 123c. 6 UAZ; 129a. 1 WNMU; 130c. 3 IPN; 131. 3 CNMA, 8 OMNH; 133. 80 LACM; 133a. 2 LACM; 133e. 2 UAZ; 133l. 9 AMNH; 135. 3 CNMA, 2 OMNH; 138. 1 OMNH; 139. 3 OMNH; 141. 2 LACM; 142. 6 CNMA, 28 OMNH, 6 UAEM; 145a. 4 UAZ; 149. 9 LACM, 2 NMNH, 1 UAZ; 149c. 1 LACM; 149e. 3 CNMA; 149f. 5 ROM; 149g. 3 NMNH; 154a. 1 OMNH; 160. 5 CNMA; 161. 5 CNMA; 167. 7 CNMA; 168. 8 LACM; 169. 7 CNMA; 170. 3 CNMA; 171. 6 CNMA, 16 OMNH; 171a. 8 CNMA, 14 OMNH; 175. 12 OMNH; 177. 1 UAEM;



178. 4 LACM; 179. 7 IPN; 180. 6 CNMA; 183. 1 IPN; 183a. 1 LSUMZ; 183b. 53 LACM; 184. 3 OMNH; 193a. 1 OMNH; 193c. 1 OMNH; 195a. 3 IPN; 198. 6 LACM; 199b. 10 LACM; 205a. 30 OMNH; 206. 2 OMNH; 210. 1 CNMA, 4 OMNH, 3 UAEM; 215. 1 LSUMZ; 216. 1 LACM, 1 UAZ; 222. 4 LACM; 224. 1 OMNH; 225. 16 OMNH; 226. 9 OMNH; 229b*. 1 CNMA. Records in literature. 8 (Kennedy et al. 1984); 12a (Handley 1966; Villa-R. 1966); 19a (Sánchez-Hernández et al. 2009), 21 (Guerrero et al. 2004); 21a (Sánchez-Hernández et al. 2009); 32b (Handley 1966; Villa-R. 1966); 43, 50, 50a, 51, 52, 52b, 52c, 53, 53a, 54, 54a, 98g (Kennedy et al. 1984); 114 (Handley 1966; Villa-R. 1966); 154a, 193b (Kennedy et al. 1984); 229* (Handley 1966); 229a* (Sánchez-Hernández et al. 1990a).

Artibeus jamaicensis

115

GREAT FRUIT-EATING BAT

Artibeus lituratus (Olfers, 1818)

Figure 6.45. Great fruit-eating bat, Artibeus lituratus.

Type locality is “Paraguay” (Olfers 1818:224) but later was restricted to “Asunción,” Paraguay, by Cabrera (1957:90). Artibeus is from the Greek arti meaning “straight, exactly fitted” and bainō meaning “to go, walk, step” (Palmer 1904; Jaeger 1955); lituratus is from the Latin, meaning “branded” or “marked,” referring to the welldefined white facial stripes of the species.

116

FAMILY PHYLLOSTOMIDAE

Description. Great fruit-eating bats are large, with a total length of 78–110 mm and forearm length greater than 67 mm (fig. 6.45). The dorsum is cinnamon-brown to dark brown, with the ventrum slightly paler. A pair of distinct whitish facial stripes is present on each side of the head; the supraorbital stripe is the most conspicuous. Ears are large. Wing membranes are blackish, and the uropatagium is narrow, concave (i.e., V-shaped), and with considerable hair on the dorsal surface. There is no external tail. The proximal onehalf of the forearm has hair. Pectoral muscles are strongly attached to skin, making specimen preparation a challenge. The cranium is robust, with postorbital processes strongly developed. Medial upper incisors are small, but larger than outer incisors, and are clearly bilobed (Davis 1984); upper incisors are separated by spaces. The m3 usually is present but not always. Dental formula is i 2/2, c 1/1, p 2/2, m 2/2 or 2/3, total 28 or 30 (Davis 1984). Distribution. Sinaloa and Tamaulipas to Paraguay and northern Argentina (Simmons 2005). Elevational range sea level to nearly 2,620 m (Muñoz Arango 1990). In Colima, 4–1,566 m. Conservation status. Least concern (IUCN 2014). No special status in Mexico (SEMARNAT 2010). Habitat. Great fruit-eating bats occupy tropical deciduous, semideciduous, and pine-oak forests. Specimens were captured in mist nets set over water and within vegetation, as well as in areas with American oil palms (Elaeis oleifera), fruiting mangos (Mangifera indica), and Madras thorns (Pithecellobium dulce). The species roosts on the underside of fronds of coconut palms (Cocos nucifera) and in other trees, as well as in caves, tree holes, and shady areas in vegetation. In Tabasco, it was captured in a cavity of a kapok tree (Ceiba pentandra; Davis 1984). Environmental information. In Colima, great fruit-eating bats were noted at 40 of 161 sites included when evaluating environmental characteristics. No significant difference was identified comparing sites where the species was recorded and not recorded for any of the 21 environmental variables. Diet. The great fruit-eating bat is frugivorous-omnivorous. In Colima, we regularly captured individuals carrying fruits of wild figs (Ficus) and nightshades (Solanum). On a coffee plantation (Coffea arabica) in Chiapas, they were closely associated with trumpet trees (Cecropia obtusifolia) and fig trees (Ficus pertusa; García-Estrada et al. 2012). In Guerrero, a captured great fruit-eating bat had the fruit of a yellow mombin (Spondias mombin) in its mouth (Lumbreras Ramos 2012) Reproduction. Examined 69 males (1 subadults, 68 adults), 84 females (2 subadults, 82 adults), and 1 adult of unknown sex.

the individual had been exposed to rabies (Lyssavirus). Measurements. Females were significantly larger than males in mass. Males were significantly larger than females in mandibular-toothrow length (appendix B). Although samples were not large, they were suggestive that total length, obtained from specimen tags, exhibited a bimodal distribution in both males and females, with modes for males at 85 and 105 mm and for females at 87 and 115 mm. The bimodal distributions are not biologically meaningful but reflect differences in how investigators measure total length in species that do not have an external tail; some measure to the posterior edge of the body of the bat not including the uropatagium, while others measure to the posterior edge of the uropatagium. Specimens examined (n = 154). 18 CNMA; 3 IPN; 36 LACM; 1 MVZ; 77 OMNH; 7 UAEM; 1 UAM-I; 11 UAZ. Localities (fig. 6.46). 3c. 5 LACM; 5a. 1 LACM; 8. 1 CNMA; 17. 2 LACM; 21. 1 OMNH; 34. 1 LACM; 43. 1 CNMA, 5 OMNH; 50. 2 OMNH; 50a. 11 OMNH; 52. 21 OMNH; 52b. 1 OMNH; 53. 7 OMNH; 53a. 4 OMNH; 68. 1 OMNH; 78. 1

Male, subadult, 1 (May). Adult, inguinal testes, 1 (January, 8×6). Scrotal testes, 15 (January, mean = 7.1×5.1 [5×3 to 10×7]). Testes (position not indicated), 3 (January, 2 [5×4, 9×5]; April, 1, [5]). Female, subadult, 2 (April, 1; May, 1). Adult, with 1 embryo, 11 (January, 4 with length only, mean = 10.7 [5, 9, 14, 15], 7 with length and width, mean = 13.0×11.3 [4×3 to 30×28]). Lactating, 4 (April, 2; May, 2). Postlactating, 3 (January, 2; June, 1). No embryo, 8 (January). The reproductive pattern of the great fruit-eating bat is continuous polyestry that is asynchronous, with 3–4 birthing periods/year (Sánchez Hernández and Romero Almaraz 1995a). Young are precocial and can fly before wing phalanges are completely ossified. Activity. We captured the species in mist nets from 1800 to 2330 h, as well as in nets in the morning when those nets were left unfurled all night. Other observations. In Colima, 2 great fruit-eating bats were tested for antirabic antibodies (Salas-Rojas et al. 2004); both tested negative. A positive test would have indicated that

Figure 6.46. Known distribution Pac ifi

Copyright © 2016, University of Oklahoma Press.

c

19°30'

O

ce

an

MEXICO

COLIMA

5

19

21

Minatitlán

19°15'

78

of great fruit-eating bat (Artibeus lituratus) in Colima.

JALISCO

83

68

79

MANZANILLO 97 98

100

104

8

COMALA

43 17 Cuauhtémoc 50 Comala VILLA DE 34 53 MINATITLÁN ÁLVAREZ 52 Villa de 122 Álvarez 149 123 CUAUHTÉMOC Colima COQUIMATLÁN Coquimatlán

133

130

COLIMA

142 145

110 Manzanillo 112

ARMERÍA 178

198

19°00'

Ixtlahuacán IXTLAHUACÁN

183 193

Great Fruit-eating Bat

3

205 Armería Tecomán 195 TECOMÁN

226

175

210

Artibeus lituratus

18°45' 20 Kilometers 10 Miles

216

MICHOACÁN

103°45'

103°30'

N 104°45'



104°30'

104°15'

104°00'

Artibeus lituratus

117

OMNH; 79. 1 OMNH; 83. 1 LACM; 97. 6 CNMA, 5 OMNH, 6 UAEM; 97c. 1 CNMA, 3 OMNH; 97d. 4 CNMA, 2 OMNH; 98g. 4 OMNH; 100. 1 CNMA, 1 OMNH; 104. 1 UAM-I; 110a. 1 UAZ; 112e. 1 UAZ; 122a. 2 LACM; 123c. 4 UAZ; 123g. 2 UAZ; 130c. 1 IPN; 133a. 2 LACM; 142. 4 CNMA, 1 UAEM; 145a. 1 UAZ; 149. 1 LACM; 149a. 1 LACM; 175. 1 OMNH; 178. 1 LACM; 183b. 13 LACM; 193a. 1 OMNH; 195. 2 IPN; 198. 6 LACM; 205a. 1 OMNH; 210. 1 OMNH; 216. 3 UAZ; 226. 3 OMNH. Records in literature. 19a, 21a (Sánchez-Hernández et al. 2009). The intermediate fruit-eating bat (Artibeus intermedius) has been considered to be a subspecies of the great

118

FAMILY PHYLLOSTOMIDAE

fruit-eating bat, but Davis (1984) concluded it represented a distinct species. Given that we do not know if specimens noted previously for Colima by Villa-R. (1966) and Kennedy et al. (1984) belong to one or the other species, we have not included the following records from the literature: 8 (Kennedy et al. 1984); 12 (Villa-R. 1966); 32b (Villa-R. 1966); 43, 50, 50a, 52, 52b, 52c, 53, 53a, 54, 54a, 98g (Kennedy et al. 1984); 133a (Villa-R. 1966); 193b (Kennedy et al. 1984). All of the numbered localities from these publications (although not necessarily the sublocalities represented by a letter) are represented by museum specimens examined and identified as great fruit-eating bats.

WRINKLE-FACED BAT

Centurio senex Gray, 1842

Figure 6.47. Wrinkle-faced bat, Centurio senex.

Type locality is “Amboyna” (Gray 1842:260), which was restricted to “Realejo, Nicaragua” by Goodwin (1946:327). Centurio is Latin meaning “centurion or commander of a company of infantry, corresponding to a captain in a modern army, whose insignia or rank is the shoulder badge or epaulet” (Palmer 1904:168); senex is Latin meaning “an old man” or “to grow old” (Jaeger 1955). The scientific name refers to the small epaulet-like tufts of white hair on the shoulders (Gray 1842; Palmer 1904), and the wrinkled face is likened to that of a “hundredyear-old-man” (Snow et al. 1980:3).



Description. The wrinkle-faced bat is small, with a total length of 55–70 mm and forearm length of 44–47 mm (fig. 6.47). The dorsum is brown or pale gray, with the venter slightly paler. White spots are present on the shoulders. The rostrum lacks hair and is flattened and rough, giving the impression of a wrinkled face. Males have well-developed ridges on the face, with skin folds on the chin; these can cover the face, as with a mask. In females, folds are smaller and the mask lacking. The noseleaf is rudimentary, and ears are large, rounded, and yellow. Striated bands are present on the membrane between the fourth and fifth digits of the wing. The uropatagium is concave (i.e., V-shaped) and lacks hair. There is no external tail. The skull has a short rostrum and high, rounded braincase with a moderately well developed sagittal crest. The palate is short, about one-half as long as wide. Upper canines have an anterior basal cavity (Hall 1981). Upper incisors are small and form a discontinuous row between the canines; inner incisors are larger than outer incisors. Lower incisors are small, short, and closely crowded, forming a deep notch in the middle (Goodwin and Greenhall 1961). Dental formula is i 2/2, c 1/1, p 2/2, m 2/2, total 28 (Goodwin and Greenhall 1961). Distribution. Sinaloa and Tamaulipas to Venezuela, and Trinidad and Tobago (Simmons 2005). Elevational range sea level to 1,450 m (Snow et al. 1980). In Colima, 4–777 m. Exhibits migration (Sanchez Hernandez 1984). Conservation status. Least concern (IUCN 2014). In Mexico, no special status (SEMARNAT 2010). Habitat. Specimens of the wrinkle-faced bat collected in January were captured in tropical deciduous forest mixed with secondary vegetation, in tropical evergreen and deciduous forests over bodies of water (Sánchez Hernández 1984; Sánchez Hernández and Romero Almaraz 1995a). The species was also netted in a lime orchard (Citrus latifolia) with a small stream running through it (Polaco and MuñizMartínez 1987) and in a deciduous forest (Watkins et al. 1972). In Nicaragua, wrinkle-faced bats were collected along a stream bordered by a bilevel gallery forest in an area mostly planted in sugar cane (Saccharum officinarum; Jones et al. 1971); some were in the lower level of the gallery forest where trees were covered with a thick network of vines. In Trinidad and Tobago, they were observed on the undersides of leaves of mango trees (Mangifera indica), dragon trees (Dracaena), and putranjivas (Putranjiva; Goodwin and Greenhall 1961). Diet. The wrinkle-faced bat is frugivorous. It consumes fruits of the azulillo tree (Vitex mollis; Ramírez-Pulido and López-Forment 1979), wild figs (Ficus), and plum trees (Prunus; Sánchez Hernández 1984). Goodwin and Greenhall (1961) reported that they suck rather than bite or crush the pulp of fruits. Reproduction. Specimens examined included 10 adult females. Female with 1 embryo, 3 (January [9×9, 10×8, 16×10]). Lactating, 3 (April). Centurio senex

119

also suggested by data for other locations in Oaxaca (2 males:9 females; Olguín Monroy et al. 2008), in Panama (0 males:22 females; Paradiso 1967), and in Trinidad (2 males:30 females; Goodwin and Greenhall 1961). Sometimes 2 wrinkle-faced bats are present in the same tree but never more than 2. Goodwin and Greenhall (1961) noted that wrinkle-faced bats cover their faces with flaps of skin from the chin. In captivity, they “slept with the skin of the chin pulled up over the face where it covered the top of the head and extended over the flaps of the ears which lie flat across the top of the head” (Goodwin and Greenhall 1961:264). Measurements. Sexual dimorphism could not be evaluated due to a lack of male specimens (appendix B). In Oaxaca, average mass of 22 males (24.5 g) and 13 females (24.1 g) was greater than that for 3 females we captured (mean 17.6 g); however, mean forearm length of 8 females (42.0 mm) from Colima was similar to means for Oaxaca (males 43.6 mm and females 43.3 mm; Santos-Moreno et al. 2010a). Specimens examined (n = 10). 4 CNMA; 2 LACM; 1 NMNH; 1 OMNH; 2 UAZ. Localities (fig. 6.48). 32a. 2 LACM; 58a. 1 NMNH; 97. 1 CNMA; 123a. 2 CNMA; 123c. 2 UAZ; 175. 1 CNMA; 226. 1 OMNH. Records in literature. 32a, 58a, 97 (Sánchez-Hernández et al. 2002); 123a (Villa-R. 1966); 226 (Sánchez-Hernández et al. 2002).

Wilson (1979) noted that reproductive data for any given region were too sparse to determine the reproductive pattern with certainty for the wrinkle-faced bat. Pregnancies in February and July on the Yucatan Peninsula suggest that the reproductive pattern for the species is bimodal polyestry. In Nayarit, 2 females caught in June contained embryos, one measuring 25 mm and the other 26 mm (Arroyo-Cabrales et al. 2008). In a mountain range in Oaxaca, wrinkle-faced bats had a bimodal reproductive pattern, with pregnant females primarily in April and August, and males with scrotal testes in June and from August through October (Santos-Moreno et al. 2010a); 9 of 10 subadults were females, as were 34 of 51 adults. In Michoacán, 6 females had embryos in February (Polaco and Muñiz-Martínez 1987), and 1 pregnant and 1 lactating female were caught in April, while a subadult was caught in November (Sanchez Hernandez et al. 1985). Overall, we estimate the reproductive pattern of the species to be bimodal polyestry. Activity. In Oaxaca, wrinkle-faced bats become active at 1800 h, with 2 activity peaks: one about midnight and another about 0300 h (Santos-Moreno et al. 2010a). One was recaptured 4.7 km from where it originally was marked (Montiel et al. 2006). Other observations. In Oaxaca, 51 wrinkle-faced bats were captured over 16 months; 17 were males and 34 were females (Santos-Moreno et al. 2010a). A female-skewed sex ratio is Figure 6.48. Known distribution

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of wrinkle-faced bat (Centurio senex) in Colima.

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VILLA DE Comala ÁLVAREZ Villa de Álvarez 123 Colima

MINATITLÁN 19°15'

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58 CUAUHTÉMOC

COQUIMATLÁN Coquimatlán

MANZANILLO

COLIMA

97 Manzanillo ARMERÍA 19°00'

Ixtlahuacán IXTLAHUACÁN Armería

Tecomán

226

175

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Wrinkle-faced Bat Centurio senex

18°45'

MICHOACÁN

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120

FAMILY PHYLLOSTOMIDAE

104°30'

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SALVIN’S BIG-EYED BAT

Chiroderma salvini Dobson, 1878

Figure 6.49. Salvin’s big-eyed bat, Chiroderma salvini.

Type locality is “Costa Rica” (Dobson 1878:533). Chiroderma is from the Greek cheir meaning “hand” and derma meaning “skin” (Jaeger 1955); salvini is a patronym honoring Osbert Salvin, a British naturalist and the collector of the type specimen.



Description. Salvin’s big-eyed bat is medium-sized, with a total length of 66–88 mm and forearm length of 40–47 mm (fig. 6.49). Dorsal color varies from pale brown to dark brown, with a narrow but prominent white median-dorsal stripe extending from the shoulders or mane to the rump. The venter is paler, varying to slightly browner or grayer. Individual hairs on the dorsum are tricolored, being fuscous, buff, and brown from bases to tips, respectively. A pair of white stripes is present, one stripe above and one below each of the large eyes. Ears are yellowish at the base and on lower margins, becoming brownish-black toward the tip; the tragus is yellowish. The noseleaf is triangular. Wing membranes that extend to the metatarsus are blackish, except for a translucent brownish area between the second and third fingers. The uropatagium, tibia, foot, and proximal two-thirds of the forearm are hairy. The uropatagium is well developed, its posterior margin being slightly concave. The tail is absent, and the feet are small. The rostrum of the skull, which lacks nasal bones, is robust and relatively narrow, and orbital ledges and processes are reduced (Handley 1966). The zygomatic arch is wide. Lower incisors are small and subequal. Inner, upper incisors are long, slender, and pointed; outer, upper incisors are small (Dobson 1878). Dental formula is i 2/2, c 1/1, p 2/2, m 2/2, total 28 (Hall and Kelson 1959; Hall 1981). Distribution. Colima, Michoacán, Hidalgo, and Chihuahua to Peru, Bolivia, and Venezuela (Simmons 2005). Elevational range sea level (Sánchez Hernández 1984) to 2,500 m (León Paniagua and Romo Vázquez 1993). In Colima, 4–1,197 m. Species migrates within the tropics (Sánchez Hernández 1984). Conservation status. Least concern (IUCN 2014). In Mexico, no special status (SEMARNAT 2010). Habitat. Salvin’s big-eyed bats were captured in mist nets set over water and in tropical deciduous forest, conifer-oak forest, secondary vegetation, and groves of fruit trees. In Guerrero, an individual was captured in gallery forest surrounded by a corn field (Zea mays) and another in tropical dry forest (Almazán-Catalán et al. 2009). In Sinaloa, they were caught in nets placed under trees with ripe fruits, such as wild figs (Ficus; Jones et al. 1972), while in Jalisco, Watkins et al. (1972) caught them in banana (Musa) groves. In Colima, they roost in caves, abandoned mines, and holes in trees. Similar observations have been reported elsewhere (Polaco and Muñiz-Martínez 1987; León Paniagua and Romo Vázquez 1993). Based on work in Costa Rica, Goodwin (1946) judged that the species probably was a tree dweller. Environmental information. Salvin’s big-eyed bats were obtained at 17 of 161 sites included in the evaluation of environmental characteristics. No significant difference was identified comparing sites where the species was recorded with sites where it was not recorded, for any of the 21 environmental variables. Chiroderma salvini

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Figure 6.50. Known distribution of Salvin’s big-eyed bat (Chiroderma salvini) in Colima.

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Cuauhtémoc 50 52 51 53 54 CUAUHTÉMOC

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100

Coquimatlán

133

COLIMA 103 Manzanillo

171

ARMERÍA

198

178 19°00'

Ixtlahuacán IXTLAHUACÁN

183 Armería

Tecomán TECOMÁN

Salvin’s Big-eyed Bat Chiroderma salvini

18°45'

MICHOACÁN

20 Kilometers 10 Miles N 104°45'

104°30'

Diet. This species is frugivorous. It consumes fruits and infrutescences, particularly those that are abundant throughout the year, such as from the trumpet tree (Cecropia obtusifolia), Mexican pepperleaf (Piper auritum), a pepper (P. pseudolindenii), giant devil’s-fig (Solanum chrysotrichum), and twoleaf nightshade (S. diphyllum; García-Estrada et al. 2012). Reproduction. Examined 14 males (1 subadult, 13 adults), 19 adult females, and 5 adults of unknown sex. Male, subadult, 1 (August). Adult, inguinal testes, 1 (January [4×3]). Scrotal testes, 5 (January, mean = 5.6×3.6 [4×2 to 7×5]). Female with 1 embryo, 3 (January, [6×5, 9, 10×6]). Lactating, 1 (April). No embryo, 1 (January). Wilson (1979) suggested that the reproductive pattern was bimodal polyestry. Data from Colima and other sources (Carter et al. 1966; LaVal 1969; Jones et al. 1972; Watkins et al. 1972; Polaco and Muñiz-Martínez 1987; GuzmánSoriano et al. 2013) indicate this probably is correct. Pregnant females were present in Colima (January [embryo, 6–10 mm), Campeche (February), Michoacán (July [18–24 mm]), Sinaloa (January [4 mm]), Jalisco (February [18–24 mm]; June [6–16 mm]), and Honduras (July [27 mm]; August [27 mm]). Lactating females were recorded in Colima (April),

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Jalisco (June), and Honduras (July, August), while in Sinaloa juveniles were recorded in May–July. Activity. Salvin’s big-eyed bats were captured in early evening from 2000 to 2200 h. In Guerrero, a bat was captured in a mist net at 2115 h (Almazán-Catalán et al. 2009), and in Sinaloa, Salvin’s big-eyed bats were in the nets by 2130 h (Jones et al. 1972). Measurements. Females were significantly larger than males in condylocanine length and mandible length (appendix B). Specimens examined (n = 38). 10 CNMA; 11 LACM; 1 NMNH; 13 OMNH; 1 TTU; 2 UAZ. Localities (fig. 6.50). 3a. 1 OMNH; 20. 1 CNMA; 21. 2 CNMA; 36. 2 LACM; 50. 1 OMNH; 50a. 1 OMNH; 51. 1 OMNH; 52. 3 OMNH; 53. 1 OMNH; 54a. 2 OMNH; 97. 3 OMNH; 97c. 1 CNMA; 100. 1 CNMA; 103. 1 CNMA; 124. 1 UAZ; 124b. 2 LACM, 1 TTU; 133a. 2 LACM, 1 NMNH, 1 UAZ; 133b. 1 CNMA; 133g. 2 CNMA; 171. 1 CNMA; 178. 1 LACM; 183b. 3 LACM; 198. 1 LACM. Records in literature. 20, 21a (Sánchez-Hernández et al. 2009); 36 (Handley 1966); 50, 50a, 52b, 53, 54a (Kennedy et al. 1984); 133g (Villa-R. 1962); 133j (Handley 1966, Villa-R. 1966, Wilson 1991); 229a* (Swanepoel and Genoways 1979).

AZTEC FRUIT-EATING BAT

Dermanura azteca (Andersen, 1906)

Figure 6.51. Aztec fruit-eating bat, Dermanura azteca.

Type locality is “Tetela del Volcan, Morelos, Mexico” (Andersen 1906:422). Dermanura is from the Greek derma meaning “skin,” anmeaning “without,” and oura meaning “tail” (Jaeger 1955); azteca refers to the Aztec people of central Mexico.



Description. The Aztec fruit-eating bat is medium-sized, with a total length of 61–71 mm and forearm length of 41–46 mm (fig. 6.51). The dorsum is blackish-brown, and the venter is paler. The face and noseleaf are broad, and ears are small. The uropatagium is narrow, with a conspicuous fringe of hair along the posterior border of the uropatagium and on its upper surface. The tail is absent. Rostrum length is slightly greater than one-half that of the braincase. Inner upper incisors are bilobed and slightly larger than outer, but not twice as large. Individuals may have a third molar, but it is reduced in size if present. Dental formula is i 2/2, c 1/1, p 2/2, m 2/2 or 2/3, total 28 or 30 (Webster and Jones 1982a). Distribution. Nuevo León and Sinaloa to Michoacán and Oaxaca to western Panama (Simmons 2005). Elevational range 575–3,000 m (Webster and Jones 1982a). In Colima, 585–1,619 m. Conservation status. Least concern (IUCN 2014; cited as Artibeus aztecus). In Mexico, no special status (SEMARNAT 2010). Habitat. One individual from January 1998 was captured in oak forest and another in a tropical semideciduous forest. The species is present in a variety of habitats. It often is associated with highlands, including cloud forests, but it can also be found in xerophytic scrub. Roosts are in mines, buildings, tunnels, and caves with high humidity. In Colima, we captured Aztec fruit-eating bats in mist nets placed across water, under wild fig trees (Ficus), and among acacia bushes (Acacia). They sometimes can be found on ventral surfaces of banana (Musa) leaves. Colonies may include dozens of individuals (Hernández-Flores and Rojas-Martínez 2010). In Chihuahua, the species was collected at the bottom of a ravine in pine-oak forest (López-González and García-Mendoza 2006). Diet. In Michoacán and Morelos, Aztec fruit-eating bats consume fruits of plants such as wild fig, capuli cherry (Prunus serotina), cypress (Cupressus), and Mexican hawthorn (Crataegus mexicana; unpublished data). Similar observations concerning their food habits were reported by Ceballos González and Galindo Leal (1984). Reproduction. Examined 2 adult males and 8 adult females. Male, scrotal testes, 1 (January [6]). Female with no embryo, 3 (January). The reproductive pattern is bimodal polyestry, with at least  2 birthing periods/year, one in February–March and the other in August–September. In Sinaloa, 18 females collected in July were pregnant (embryos 19–26 mm; Jones et al. 1972), while in the state of México, a young female bat was being carried by its mother in September (Villa-R. 1966).

Dermanura azteca

123

In Chihuahua, a lactating female was captured at the end of August (López-González and García-Mendoza 2006). Young are precocial. Activity. Aztec fruit-eating bats were captured early in the evening with activity diminishing by 2200 h. They became active again in early morning. Other observations. Simmons (2005) did not recognize Dermanura as generically distinct from Artibeus. We follow Owen (1987) and a number of subsequent authors (e.g., Baker et al. 2003; Ramírez-Pulido et al. 2014) in recognizing the taxonomic validity of Dermanura.

Measurements. Males were significantly larger than females in total length, ear length, and mass (appendix B). Specimens examined (n = 10). 5 CNMA; 1 LACM; 4 OMNH. Localities (fig. 6.52). 1e. 1 OMNH; 13. 1 OMNH; 19. 1 CNMA; 32a. 1 LACM; 49. 3 CNMA; 52. 1 OMNH; 54a. 1 CNMA, 1 OMNH. Records in literature. 1e, 13a (Sánchez-Hernández et al. 2002); 13c (Kennedy et al. 1984); 19 (Sánchez-Hernández et al. 2002); 19a (Sánchez-Hernández et al. 2009); 32a (Sánchez-Hernández et al. 2002); 54a (Kennedy et al. 1984; Sánchez-Hernández et al. 2002).

Figure 6.52. Known distribution

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MINATITLÁN 19°15'

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Cuauhtémoc 52 54 CUAUHTÉMOC

COQUIMATLÁN Coquimatlán

MANZANILLO

COLIMA Manzanillo ARMERÍA 19°00'

Ixtlahuacán IXTLAHUACÁN Armería

Tecomán TECOMÁN

Aztec Fruit-eating Bat Dermanura azteca

18°45'

MICHOACÁN

20 Kilometers 10 Miles N 104°45'

124

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104°30'

104°15'

104°00'

103°45'

103°30'

PYGMY FRUIT-EATING BAT

Dermanura phaeotis Miller, 1902

Figure 6.53. Pygmy fruit-eating bat, Dermanura phaeotis.

Type locality is “Chichen Itza, Yucatan” (Miller 1902:405). Dermanura is from the Greek derma meaning “skin,” an- meaning “without,” and oura meaning “tail” (Jaeger 1955); phaeotis is of Greek derivation, coming from phaios meaning “dusky” and referring to the dusky gray coloration of the species (Timm 1985).



Description. The pygmy fruit-eating bat is small, with a total length of 51–60 mm and forearm length of 35–42 mm (fig. 6.53). Dorsal coloration is uniform, varying from grayish-brown to dark brown. Hair is moderately thick and smooth. The basal one-half of the forearm, both dorsally and ventrally, is well furred. Wing membranes are blackish, except for the tips, where pigment is lacking. The noseleaf is thick and pointed, being longer than wide, while ears are rounded and generally with a yellowish fringe. Whitish facial stripes are present and well developed above and below the eye. The uropatagium is narrow, concave, and without hair along the posterior border. The rostrum is short, broad, and up-turned. The M1 has a relatively broad talon, composing one-fourth of the total breadth of toothrow. The third molar is absent. Dental formula is i 2/2, c 1/1, p 2/2, m 2/2, total 28 (Timm 1985). Distribution. Veracruz, Sinaloa, and Michoacán to Ecuador and Guyana (Simmons 2005). Elevational range sea level to 1,216 m (Hall 1981). In Colima, 4–1,566 m. Apparently, more common at lower elevations. Exhibits local movements (Sánchez Hernández 1984). Conservation status. Least concern (IUCN 2014; cited as Artibeus phaeotis). In Mexico, no special status (SEMARNAT 2010). Habitat. The pygmy fruit-eating bat occurs in various habitats, such as deciduous forests, evergreen tropical forests, orchards, and secondary vegetation. We captured it in mist nets set in vegetation and across water. They roost in palm fronds, under leaves of banana trees (Musa), in tree holes, and in shady areas of trees. In Tabasco, they were caught in nets placed among cacao trees (Theobroma cacao) and coconut palms (Cocos nucifera; Sánchez Hernández and Romero Almaraz 1995a). In Costa Rica, Timm (1985) noted that individuals modified banana leaves to form a tent where they roosted during the day. Environmental information. Pygmy fruit-eating bats were documented at 71 of the 161 localities used in the evaluation of 21 environmental characteristics. It tended to be at sites with, on average, higher precipitation seasonality (111.1% at sites where recorded vs. 109.5% where not recorded). Thus, the variation among monthly precipitation values over the 12-month period was greater where the species was captured. Diet. Pygmy fruit-eating bats are frugivorous-omnivorous. They consume the fruit of wild figs (Ficus), a genus of Moraceae (Trophis), and nightshades (Solanum; Sánchez Hernández 1984). On a coffee (Coffea arabica) plantation in Chiapas, they fed on fruits and frutescences of 13 species, the most important being the giant devil’s-fig (Solanum chrysotrichum) and trumpet tree (Cecropia obtusifolia; García-Estrada et al. 2012). They also consume pollen and insects (Timm 1985). Dermanura phaeotis

125

Reproduction. Examined 234 males (9 subadults, 225 adults), 187 females (3 subadults, 184 adults), and 2 adults of unknown sex. Male, subadult, 9 (March, 2; April, 1; May, 4; July, 2). Adult, abdominal testes, 6 (January, 5, mean = 4.7×3.0 [3×2 to 6×4]; September, 1 [8×5]). Inguinal testes, 13 (January, mean = 4.4×2.8 [1×1 to 7×5]). Scrotal testes, 94 (January, mean = 5.0×3.4 [3×2 to 11×5]). Testes (position not indicated), 1 (January [4×3]). Female, subadult, 3 (March, 2; April, 1). Adult with 1 embryo, 60 (January, 19 with length only, mean = 15.2 [5 to 23], 41 with length and width, mean = 15.9×10.8 [1×1 to 25×17]). Lactating, 19 (March, 4; April, 1; June, 2; July, 6; August, 4; October, 2). Postlactating, 1 (January). Not pregnant, 19 (January, 14; August, 1; October, 4). The reproductive pattern of the pygmy fruit-eating bat is continuous polyestry, with at least 3 asynchronous birthing periods/year. It has postpartum estrus (Sánchez Hernández and Romero Almaraz 1995a). Young are precocial. In Jalisco, pregnant females were caught in January, April, and June, and a lactating female was caught in August (Watkins et al. 1972). In Sinaloa, 4 pregnant females were caught in July (embryos 21–28 mm; Jones et al. 1972), while in Tabasco, there were pregnant or lactating females in January, March,

September, and November (Sánchez Hernández and Romero Almaraz 1995a). In Michoacán, 13 females in total were captured in March–May and December, and individual lactating females were caught in February, March, and May (Sanchez Hernandez et al. 1985). Activity. Pygmy fruit-eating bats were captured from 1900 to 2200 h. Their active period begins about 1830 h and diminishes by 2100–2200 h (Sánchez Hernández and Romero Almaraz 1995a). In Campeche, some were recaptured up to 4.7 km from where they were originally marked (mean 1.8 km; Montiel et al. 2006). Other observations. In Colima, 6 pygmy fruit-eating bats were evaluated for the presence of antirabic antibodies (Salas-Rojas et al. 2004); a positive test indicates an individual had been exposed to rabies (Lyssavirus). Of the 6 analyzed, 3 from a disturbed area (agricultural area) tested negative, while 3 from an undisturbed area (tropical semideciduous forest) tested positive. Systematic relationships of smaller bats of the genus Artibeus (in which this species is sometimes placed) have been an issue of contention; refer to comments regarding validity of the genus Dermanura under the account for the Aztec fruit-eating bat (Dermanura azteca).

Figure 6.54. Known distribution JALISCO

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40 43 17 Cuauhtémoc41 42 52 50 Comala 33 32 49 VILLA DE 48 53 34 ÁLVAREZ 122 54 Villa de 57 58 Álvarez 149 123 CUAUHTÉMOC Colima 154 COQUIMATLÁN

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65 81

97 103

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MANZANILLO

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109 Manzanillo 114 113 112

Coquimatlán 133 140 141 142 145

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183 184

187

Pygmy Fruit-eating Bat

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167

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ARMERÍA 180

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COLIMA

192

Armería

Tecomán

206

226

TECOMÁN

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Dermanura phaeotis

18°45'

214 215

20 Kilometers 10 Miles

MICHOACÁN

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104°30'

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Measurements. Females were significantly larger than males in 9 of 16 measurements: mass, forearm length, third-metacarpal length, greatest skull length, condylocanine length, maxillary-toothrow length, interorbital constriction, braincase breadth, and mandibular-toothrow length (appendix B). Total length, garnered from specimen tags, exhibited a bimodal distribution in both males and females, with modes for males at 55 and 65 mm and for females at 55 and 67 mm. The bimodal distributions are not biologically meaningful but reflect differences in how investigators measure total length in species that do not have an external tail; some measure to the posterior edge of the body of the bat not including the uropatagium, while others measure to the posterior edge of the uropatagium. Specimens examined (n = 421). 2 AMNH; 118 CNMA; 2 FMNH; 16 IPN; 1 KU; 67 LACM; 5 LSUMZ; 1 MVZ; 14 NMNH; 134 OMNH; 3 ROM; 16 UAEM; 40 UAZ; 2 WNMU. Localities (fig. 6.54). 8a. 2 WNMU; 12a. 5 LACM; 12f. 1 CNMA; 17. 2 LACM; 21. 5 CNMA, 5 OMNH, 2 UAEM; 32a. 4 LACM; 32b. 8 CNMA; 32e. 16 LACM; 32j. 1 LACM; 33b. 4 UAZ; 34. 1 LACM; 40. 2 OMNH; 41. 1 OMNH; 42. 10 CNMA, 4 OMNH, 1 UAEM; 43. 4 OMNH; 48. 1 OMNH; 49. 2 OMNH; 50a. 5 OMNH; 52. 16 OMNH; 52b. 1 OMNH; 53. 5 OMNH; 53a. 2 OMNH; 54. 1 OMNH; 57. 12 LACM; 60. 2 OMNH; 60a. 1 CNMA, 3 OMNH, 2 UAEM; 61. 1 UAEM; 62. 6 LACM; 65. 4 OMNH; 67. 1 CNMA, 3 OMNH, 2 UAEM; 68. 5 CNMA, 8 OMNH, 1 UAEM; 70. 3 CNMA, 5 OMNH, 1 UAEM; 72. 3 OMNH; 72a. 4 OMNH, 1 UAEM; 73. 1 OMNH;



75. 1 CNMA, 1 OMNH; 76. 1 UAZ; 77. 1 CNMA, 2 OMNH; 81. 1 OMNH; 82. 1 LACM; 85. 2 LACM; 88a. 1 CNMA; 88b. 1 OMNH; 90. 1 CNMA; 91. 2 CNMA; 92. 1 CNMA, 1 OMNH; 97. 3 CNMA, 1 OMNH; 97a. 1 OMNH; 97c. 1 CNMA; 103. 3 CNMA, 5 OMNH; 109. 1 KU; 112b. 1 UAZ; 112c. 1 UAZ; 112e. 2 UAZ; 113. 1 CNMA, 20 OMNH; 114f. 2 CNMA; 114g. 3 CNMA; 114l. 1 MVZ; 115. 1 CNMA; 116. 3 CNMA; 116a. 2 CNMA; 116b. 4 CNMA, 12 OMNH; 122a. 3 LACM; 123a. 6 CNMA; 123c. 5 UAZ; 123g. 2 UAZ; 130b. 1 AMNH, 9 CNMA; 130c. 7 IPN; 131. 2 CNMA, 2 OMNH; 133. 4 LACM; 133a. 1 AMNH, 3 UAZ; 133b. 14 CNMA; 133e. 16 UAZ; 133l. 2 FMNH, 13 NMNH; 140. 1 CNMA; 142. 1 OMNH, 5 UAEM; 145a. 1 UAZ; 149. 3 LACM; 149f. 3 ROM; 149g. 1 NMNH; 161a. 1 UAZ; 167. 2 CNMA; 171. 1 OMNH; 171a. 2 CNMA; 179. 9 IPN; 180. 17 CNMA; 183b. 3 LACM; 184a. 1 CNMA; 187c. 3 UAZ; 192. 2 LACM; 199b. 1 LACM; 206. 1 OMNH; 214. 2 LSUMZ; 215. 3 LACM; 222. 1 LACM; 225. 1 OMNH; 226. 1 OMNH. Records in literature. 8 (Kennedy et al. 1984); 12 (Villa-R. 1966, Davis 1970); 21a (Sánchez-Hernández et al. 2009); 34a (Davis 1970); 43, 50, 50a, 52, 52b, 53, 53a, 54 (Kennedy et al. 1984); 58c, 76, 112e (Davis 1970); 114, 114g (Villa-R. 1966); 122 (Davis 1970); 123a (Villa-R. 1966); 123c (Davis 1970); 130, 133a (Villa-R. 1966), 133i (Davis 1970); 133l (Andersen 1908; Davis 1958); 141, 145, 149a (Davis 1970); 154a (Kennedy et al. 1984); 161a (Davis 1970); 184a (Villa-R. 1966); 187c, 199a, 214, 215a (Davis 1970); 229a* (Andersen 1906); 233a** (Villa-R. 1966); 234** (Davis 1970).

Dermanura phaeotis

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TOLTEC FRUIT-EATING BAT

Dermanura tolteca (de Saussure, 1860)

Figure 6.55. Toltec fruit-eating bat, Dermanura tolteca.

Type locality is “Mexique” (de Saussure 1860a:425), subsequently restricted to “Mirador, Veracruz” (Hershkovitz 1949:449). Dermanura is from the Greek derma meaning “skin,” anmeaning “without,” and oura meaning “tail” (Jaeger 1955); tolteca is in reference to the Toltec culture, an archaeological Mesoamerican culture of Mexico.

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Description. Toltec fruit-eating bats are small, with a total length of 51–63 mm and forearm length of 38–42 mm (fig. 6.55). Dorsal coloration in adults varies from pale brown to dark brown. Color variation is probably related to several factors, including latitude, age, reproductive condition, and humidity at roosting sites. Facial stripes are present but diffuse. Ears have a yellowish fringe and are not bordered with fur (Villa-R. 1966). The uropatagium is narrow, concave, and conspicuously bordered by hair of variable length. On the cranium, anterior portions of nasal bones are nearly horizontal (slightly elevated). Dental formula is i 2/2, c 1/1, p 2/2, m 2/2, total 28 (Webster and Jones 1982b). Distribution. Northern Sinaloa, central Nuevo León, and Tamaulipas to northeastern Colombia and Ecuador (Webster and Jones 1982b). Elevational range sea level (Sánchez Hernández 1984) to about 2,500 m (León Paniagua and Romo Vázquez 1993). In Colima, 4–2,220 m. Most widely distributed below 300 m but also captured at about 1,220 m (Jones et al. 1972). Undertakes local migrations (Sánchez Hernández 1984). Conservation status. Least concern (IUCN 2014; cited as Artibeus toltecus). In Mexico, no special status (SEMARNAT 2010). Habitat. The Toltec fruit-eating bat occupies tropical deciduous forests with mesquites (Prosopis) and prickly pears (Opuntia); they also dwell in xerophytic scrub and conifer-oak forest. This species was captured in mist nets set in vegetation with American oil palms (Elaeis oleifera), mango trees (Mangifera indica), and wild fig trees (Ficus) in addition to along banks of water bodies (Webster and Jones 1982b). They roost on the underside of palm fronds (Sánchez Hernández 1984). Toltec fruit-eating bats roost at sites where shadows or darkness are total or near total, including in caves, abandoned mines, and hollow trees, as well as in tents formed from leaves of banana trees (Musa) and coconut palms (Cocos nucifera). In Oaxaca, they were captured in a coniferous forest (Alvarez and Ramírez-Pulido 1972) while in Chihuahua, they were caught along a creek at the bottom of an open valley surrounded by tropical deciduous forest (López-González and García-Mendoza 2006). Environmental information. In Colima, Toltec fruit-eating bats were at 57 of 161 sites used to evaluate 21 environmental characteristics. The species tended to be at higher elevations (mean elevation of 618 m where species was recorded vs. 335 m where not recorded) and at sites with higher annual precipitation (1,008.3 vs. 938.0 mm), mean diurnal temperature range (13.4 vs. 13.1°C), and temperature annual range (19.1 vs. 18.3°C), but lower temperature seasonality (15.1 vs. 15.7°C). Locations where they occurred also tended to have more precipitation in coldest quarter (38.6 vs. 34.6 mm), wettest month (252.6 vs. 237.9 mm), and wettest quarter

10×5]; February, 1 [7×5]; July, 2 [5×4]; October, 1 [nm]). Testes (position not indicated), 9 (January, 4, mean = 4.0×3.0 [3×2, 4×3, 4×4, 5×3]; April, 5, mean = 5.2 [4 to 6]). Female, subadult, 2 (May, 1; June, 1). Adult with 1 embryo, 47 (January, 18 with length only, mean = 7.2 [3 to 14], 21 with length and width, mean = 8.1×6.4 [3×3 to 17×15], 2 [nm]; April, 1 [4]; July, 2 [5×4, nm]; October, 1 [5×3]; November, 1 [6×4]; December, 1 [11×9]). With 2 embryos, 1 (July). Lactating, 17 (May, 5; August, 11; September, 1). Lactating with 1 embryo, 1 (April [8]). Postlactating, 4 (January, 1; May, 2; June, 1). Not pregnant, 27 (January, 23; April, 2; September, 1; October, 1). The reproductive pattern is continuous polyestry with postpartum estrus. It is possible that there are 3 birthing periods/year. A single young is born after a 3-month gestation; young are precocial, born covered with hair and with eyes open. Lactation lasts less than 1 month. Wilson (1979) reported pregnant females in each month from January– August and in October. In Michoacán, Sanchez Hernandez et al. (1985) recorded 2 pregnant females in September, as well as 1 pregnant and lactating in March. In Sinaloa, pregnant females were captured in January, May, and October, with lactating females present in May (Jones et al. 1972).

(694.1 vs. 638.1 mm). Those sites tended to have lower annual mean temperature (24.3 vs. 25.5°C), isothermality (69.9 vs. 71.0%), and maximum temperature in warmest month (33.1 vs. 33.7°C), as well as lower mean temperature in coldest quarter (22.1 vs. 23.3°C), driest quarter (23.0 vs. 23.9°C), warmest quarter (26.0 vs. 27.2°C), and wettest quarter (25.4 vs. 26.8°C). Minimum temperature in coldest month was also lower (14.0 vs. 15.4°C). Diet. Fruits are major dietary items for Toltec fruit-eating bats, especially wild figs (Ficus) and cecropias (Cecropia; Gardner 1977). In Chiapas on a coffee (Coffea arabica) plantation, they fed on fruits and frutescences of 15 species (García-Estrada et al. 2012), the most prominent being giant devil’s-fig (Solanum chrysotrichum), twoleaf nightshade (S. diphyllum), trumpet tree (Cecropia obtusifolia), and salvia (Salvia). Reproduction. Examined 199 males (2 subadults, 197 adults), 231 females (2 subadults, 229 adults), and 5 adults of unknown sex. Male, subadult, 2 (May). Adult, abdominal testes, 10 (January, mean = 4.7×3.2 [4×2 to 7×4]). Inguinal testes, 1 (January [1×1]). Scrotal testes, 68 (January, 20 with length only, mean = 4.9 [4 to 8], 44 with length and width, mean = 5.2×3.7 [3×2 to

Figure 6.56. Known distribution Pac ifi

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COMALA 27 44 40 41 17 Cuauhtémoc 42 30 52 50 47 Comala 32 49 VILLA DE 48 53 MINATITLÁN 55 122 35 36 ÁLVAREZ Villa de 54 57 58 59 Álvarez CUAUHTÉMOC Colima 154 COQUIMATLÁN 29

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ARMERÍA 179

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Ixtlahuacán IXTLAHUACÁN

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Toltec Fruit-eating Bat

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Dermanura tolteca

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In Jalisco, Watkins et al. (1972) caught pregnant females in January–April and June–July, while lactating females were recorded in January, June–July, and September. Volant young were recorded in August–September in Jalisco (Watkins et al. 1972). Activity. Toltec fruit-eating bats typically exit shelter before nightfall, flying within vegetation, over treetops, or over streams and riverbeds. Most captures in Colima were from 1900 to 2200 h. In Veracruz, average distance between recapture sites for 4 individuals was 295 m (0–417 m), with recaptures occurring 30–122 days after initial capture (Galindo-González and Sosa 2003). Flight is slow and delicate, somewhat like that of a butterfly. Other observations. In Colima, 17 Toltec fruit-eating bats were evaluated for presence of antirabic antibodies (Salas-Rojas et al. 2004); a positive test indicates an individual had been exposed to rabies (Lyssavirus). Of 15 from a disturbed area (agricultural area), 1 tested positive, and the 2 from an undisturbed area (tropical semideciduous forest) both tested positive. Systematic relationships of the smaller bats of the genus Artibeus (in which this species is sometimes placed) have been an issue of contention; refer to comments regarding the validity of the genus Dermanura under the account for the Aztec fruit-eating bat (Dermanura azteca). Measurements. Females were significantly larger than males in mass, forearm length, third-metacarpal length, condylocanine length, interorbital constriction, and mandible length (appendix B). Total length, obtained from specimen tags, exhibited a bimodal distribution in both males and females, with modes for males at 58 and 68 mm and for females at 58 and 67 mm. The bimodal distributions are not biologically meaningful but reflect differences in how investigators measure total length in species that do not have

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an external tail; some measure to the posterior edge of the body of the bat not including the uropatagium, while others measure to the posterior edge of the uropatagium. Specimens examined (n = 435). 3 AMNH; 72 CNMA; 13 IPN; 115 LACM; 3 MVZ; 212 OMNH; 15 UAEM; 2 UAZ. Localities (fig. 6.56). 1. 2 OMNH; 1d. 1 CNMA, 4 OMNH; 1e. 1 OMNH; 1f. 2 CNMA; 2. 2 CNMA, 2 UAEM; 3. 1 OMNH; 3a. 6 CNMA, 4 OMNH, 2 UAEM; 3b. 10 LACM; 8. 18 OMNH; 12a. 1 LACM; 17. 1 LACM; 21. 13 CNMA, 15 OMNH, 5 UAEM; 27a. 3 IPN; 29. 1 LACM; 30. 2 LACM; 32a. 2 LACM; 32b. 1 CNMA; 32j. 1 LACM; 35. 1 LACM; 36. 2 LACM; 40. 4 OMNH; 40a. 2 OMNH; 41. 1 CNMA, 5 OMNH; 42. 25 CNMA, 4 UAEM; 44a. 2 OMNH; 47. 1 CNMA; 48. 1 OMNH; 49. 1 CNMA, 1 OMNH; 50. 2 OMNH; 50a. 19 OMNH; 52. 2 CNMA, 47 OMNH; 52a. 2 OMNH; 52b. 1 OMNH; 53. 37 OMNH; 53a. 3 OMNH; 54a. 1 CNMA, 1 OMNH; 55. 2 OMNH; 57. 3 LACM; 59. 1 CNMA; 65. 1 OMNH; 67. 1 OMNH; 68. 7 OMNH; 70. 1 OMNH; 81. 2 CNMA; 88. 1 OMNH; 88a. 1 OMNH; 88b. 1 OMNH; 89. 1 OMNH; 90. 2 OMNH; 91. 1 OMNH; 92. 1 OMNH; 97. 1 CNMA, 1 OMNH, 1 UAEM; 97b. 1 CNMA; 98g. 1 OMNH; 101. 1 OMNH; 110a. 3 MVZ; 112c. 1 UAZ; 113. 1 OMNH; 122a. 8 LACM; 130b. 2 CNMA; 130c. 2 IPN; 130f. 1 AMNH; 131. 2 CNMA, 2 OMNH; 133. 1 LACM; 133a. 2 AMNH; 137. 1 OMNH; 154a. 2 OMNH; 158. 4 CNMA; 179. 8 IPN; 180. 1 CNMA; 183b. 60 LACM; 187a. 1 LACM; 187d. 1 UAZ; 192. 3 LACM; 198. 17 LACM; 205a. 2 CNMA, 10 OMNH, 1 UAEM; 216. 1 LACM; 229b*. 1 OMNH. Records in literature. 5d (Tellez-Giron et al. 1997); 8 (Kennedy et al. 1984); 11c, 12 (Davis 1969); 21a (Sánchez-Hernández et al. 2009); 50, 50a, 52b, 53, 53a, 54a (Kennedy et al. 1984); 58b (Davis 1969); 98g (Kennedy et al. 1984); 112c (Davis 1969); 122 (Davis 1969); 130 (Villa-R. 1966); 154a (Kennedy et al. 1984); 187b, 216 (Davis 1969).

VELVETY FRUIT-EATING BAT

Enchisthenes hartii (Thomas, 1892)

Figure 6.57. Velvety fruit-eating bat, Enchisthenes hartii.

Type locality is “Trinidad” (Thomas 1892:408), which later was restricted to “Botanic Gardens,” Port-of-Spain, Trinidad (Thomas 1893:163). Enchisthenes is based on Greek words enchos meaning “spear” and sthenos meaning “strength,” indicating “armed with a spear” in reference to the form of the erect portion of the noseleaf (Andersen 1906); hartii honors J. H. Hart, superintendent of the Botanic Gardens in Portof-Spain, Trinidad. Hart presented a collection of bats to the British Museum, including the type specimen (Thomas 1892).



Description. The velvety fruit-eating bat (fig. 6.57) is small, with a total length of 60 mm and forearm length of 37–43 mm (Arroyo-Cabrales and Owen 1997). The dorsum is chocolate-brown, with the venter paler. The face is short and ears small; the tragus has a pointed projection on the inner margin, about 1 mm below the tip. The noseleaf is medium in size and wide. Facial stripes, narrow and buffy, may or may not be present. The uropatagium is narrow, with hair along the posterior margin. The proximal one-half of the forearm and lower leg are densely furred. The skull is moderately wide. Medial upper incisors are unicuspid. The M3 is in a row situated directly behind M2 and is as broad as the posterior margin of M2; m3 is about one-fourth the size of m2. The well-developed M3/m3 affects the form of the surrounding bone (Arroyo-Cabrales and Owen 1996). Dental formula is i 2/2, c 1/1, p 2/2, m 3/3, total 32 (Miller 1907). Distribution. Michoacán, Jalisco, and Tamaulipas to Bolivia and Venezuela, plus Trinidad (Simmons 2005). Elevational range sea level to 3,500 m (Arroyo-Cabrales and Owen 1997). In Colima, 1,258 m. Conservation status. Least concern (IUCN 2014). In Mexico, protected (SEMARNAT 2010). Habitat. The velvety fruit-eating bat occurs in several habitats, including tropical deciduous, semideciduous, evergreen, and pine-oak forests. Roosting sites are unknown, but it is assumed that they roost in caves, abandoned mines, and holes in trees. They have been captured in mist nets set over water, in gallery forest, and among trees of common guava (Psidium guajava) and banana (Musa; Almazán-Catalán et al. 2009), as well as wild fig trees (Ficus) with fruit (Villa-R. 1966). Diet. The velvety fruit-eating bat is frugivorous and consumes wild figs (Arroyo-Cabrales and Owen 1997). In Jalisco, velvety fruit-eating bats were common in groves of wild figs where they were eating small fruits about 1 cm in diameter; they quickly snipped the fruits from a tree while in flight and carried them to a convenient perch to be eaten (de la Torre 1955). Reproduction. Examined 1 adult female. Lactating with 1 embryo, 1 (April [6]). The reproductive pattern is bimodal polyestry. Gardner et al. (1970) suggested they are reproductively active throughout the year, but data from several sources (i.e., Honduras, LaVal 1969; Costa Rica, Gardner et al. 1970; Chiapas, Baker et al. 1971; Colombia, Arroyo-Cabrales and Owen 1997; El Salvador, Hellebuyck et al. 1985; Colima, herein), albeit somewhat limited, suggest that there may be a bimodal pattern. Pregnant females are known during January–February and May–June, while lactating females have been recorded in May–August. Females, both lactating and pregnant, have been recorded in April and August.

Enchisthenes hartii

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Figure 6.58. Known distribution JALISCO

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Ixtlahuacán IXTLAHUACÁN

Velvety Fruit-eating Bat

Armería

Tecomán TECOMÁN

Enchisthenes hartii

18°45'

MICHOACÁN

20 Kilometers 10 Miles N 104°45'

104°30'

In Honduras, 1 lactating female with a small embryo was caught in August (LaVal 1969). In Michoacán, 1 lactating female was recorded in April (Sanchez Hernandez et al. 1985), while in Guerrero, 2 lactating females were obtained in August (Almazán-Catalán et al. 2009). Activity. In Guerrero, the species was captured at 2200 h (Almazán-Catalán et al. 2009).

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Measurements. Sexual dimorphism was not analyzed due to there being only a single specimen (appendix B). Specimens examined (n = 1). 1 LACM. Localities (fig. 6.58). 5a. 1 LACM. Records in literature. 5b (Arroyo-Cabrales and Owen 1996).

Family Mormoopidae (Mustached Bats) The family contains 2 genera and 10 species (Simmons 2005). Mustached bats are insectivores that occupy arid, subtropical, and tropical habitats from the southwestern United States, across most of Mexico and Central America, and into southeastern Brazil. They are most common in tropical forests, but some large colonies are present in desert habitats. Mormoopids have a conspicuous, leaflike flap of skin on the lower lip, and the mouth is surrounded by short, bristlelike hairs; these structures may aid in echolocation, respiration, capturing insects, or a combination of these functions. Size varies from 7 to 20 g, wingspan ranges from 28 to 39 cm, and color of pelage may be cinnamon with various shades of grays and browns. Mustached bats usually are colonial, and they inhabit caves, tunnels, and abandoned buildings where they are able to crawl on walls with considerable agility. Day roosts, which may contain thousands of individuals, usually are in the hottest and most humid parts of tropical caves and abandoned mines. Mustached bats from large colonies may disperse many kilometers from day-roosting sites to forage at night. Beneath day roosts, the accumulation of nitrogen-rich guano is often harvested by humans for use as fertilizer (Nowak 1999; Harvey et al. 2011; Vaughan et al. 2015). For Colima, 1,034 museum specimens document 2 genera and 4 species as occurring in the state, including Peters’s ghost-faced bat (Mormoops megalophylla, 160 specimens), Davy’s naked-backed bat (Pteronotus davyi, 201), common mustached bat (P. parnellii, 374), and Wagner’s mustached bat (P. personatus, 299). A mixed colony of common mustached bats and Wagner’s mustached bats in a cave is shown in figure 6.59.

Figure 6.59. Family Mormoopidae. Common mustached bats (Pteronotus parnellii) and Wagner’s mustached bats (Pteronotus personatus) in a mixed colony in a cave.

133

PETERS’S GHOST-FACED BAT

Mormoops megalophylla (Peters, 1864)

Figure 6.60. Peters’s ghost-faced bat, Mormoops megalophylla.

Type locality is “Mexico” (Peters 1864:382), which was restricted to “Parrás [Parras], Coahuila, México” by Smith (1972:116). Mormoops is from the Greek mormo meaning “a bugbear” and combined with the Greek ops indicating “appearance” (Jaeger 1955); megalophylla is from the Greek megas meaning “great” and phyllon meaning “leaf” (Rezsutek and Cameron 1993).

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Description. Peters’s ghost-faced bat is medium-sized, with a total length of 65–75 mm and forearm length of more than 50 mm (fig. 6.60). The dorsum is reddish-brown or brown, with long, smooth, and silky pelage. On young, the dorsum is pale brown. The lower lip and chin exhibit epidermal folds. Each nostril is surrounded by an individual pad; pads are separated by a long ridge with wart-like tubercles. Eyes are small, and ears large, rounded, and united by a membranous band. The tragus is complex, with a prominent secondary fold. The uropatagium and wing membrane attach to the ankle by a short ligament not bound to the tibia. About one-half of the tail protrudes from the dorsum of the uropatagium. The skull has a raised rostrum and globose braincase, and is so greatly shortened that both the rostrum and braincase are broader than long. The crown of the head is so greatly elevated above the face line that the foramen magnum is above the level of the rostrum. Lower incisors are about equal in size and in a continuous row; they are trilobate and crowded between canines. Dental formula is i 2/2, c 1/1, p 2/3, m 3/3, total 34 (Goodwin 1946; Hall 1981; Rezsutek and Cameron 1993). Distribution. Southern Arizona and Texas in the United States and from Baja California Peninsula to Honduras (not recorded from Honduras to Isthmus of Panama); in South America along the Caribbean coast of Colombia and Venezuela, the Netherlands Antilles and Trinidad; and along Pacific coasts of Colombia and Ecuador (Rezsutek and Cameron 1993). Elevational range sea level to more than 3,000 m (Rezsutek and Cameron 1993). In Colima, 4–1,619 m. Exhibits migratory movements dependent on geographical location. For example, in Tío Bartolo Cave in Nuevo León, the colony had more than 500,000 individuals in November but 50,000 in December; the cave was occupied by Brazilian free-tailed bats (Tadarida brasiliensis) in late January. In Morelos, largest numbers were encountered in September (Villa-R. 1966). Conservation status. Least concern (IUCN 2014). In Mexico, no special status (SEMARNAT 2010). Habitat. Peters’s ghost-faced bat was captured among vegetation or along sides of rivers and streams. When caught in caves, they were lethargic and slow to become active. A colony of several thousand Peters’s ghost-faced bats was 20 m wide in a cave where temperature and humidity were high. The species was also found in abandoned mines and in tunnels where temperature and humidity were high and many insects were present (Sánchez Hernández 1984). Some were captured in mist nets 5 m above ground among mangos (Mangifera indica), wild figs (Ficus), guavas (Psidium), Madras thorns (Pithecellobium dulce), and acacias (Acacia). Peters’s ghost-faced bats inhabit tropical semideciduous and deciduous forests, xerophytic shrublands, and relict patches of coniferous and oak forests. In Sinaloa, a colony included 400,000–800,000 bats, of which about 4% were Peters’s

ghost-faced bats (Bateman and Vaughan 1974). In Chihuahua, they roosted inside a mine that also contained Mexican greater funnel-eared bats (Natalus mexicanus), Californian leaf-nosed bats (Macrotus californicus), and common mustached bats (Pteronotus parnellii; López-González and García-Mendoza 2006). Roosting places of Peters’s ghost-faced bats within 4 caves in central Mexico had a mean temperature of 32.9°C (range 16.2–37.0°C) and mean humidity of 64.2% (46.6–100%; Avila-Flores and Medellín 2004). Caves were intermediate to large in size and in limestone or clastic rock. Environmental information. In Colima, Peters’s ghost-faced bats were recorded at 29 of 161 sites used when evaluating environmental characteristics. No significant difference was identified when comparing sites where the species was recorded and not recorded for any of the 21 environmental variables. Diet. Peters’s ghost-faced bats are insectivorous. They feed on large-bodied moths (Lepidoptera; Easterla and Whitaker 1972). In Guerrero, stomach contents by volume included mainly Lepidoptera but also some Coleoptera and Trichoptera (Sil Berra 2010). Reproduction. Examined 81 adult males, 78 adult females, and 1 adult of unknown sex.

Male, abdominal testes, 5 (January, mean = 3.7×2.5 [2×1 to 6×4]). Inguinal testes, 3 (January [1×1, 2×1, 4×2]). Scrotal testes, 7 (January, mean = 3.0×2.3 [2×2 to 5×3]). Testes (position not indicated), 10 (January, mean = 2.9×1.5 [1×1 to 7×3]). Female with no embryo, 30 (January, 20; March, 10). In Colima, 8 pregnant females were captured in May (López Wilchis and Torres Flores 2005). Females with an embryo have been recorded as follows: Michoacán, 1, May (Sanchez Hernandez et al. 1985); Campeche, 2, February (Jones et al. 1973); Arizona, 1, June (Barbour and Davis 1969); Coahuila, 1 (18 mm), March; Nuevo León, 3, May; and Sonora, 8 in April, 1 in May (8–23 mm; Villa-R. 1966). Apparently, the reproductive pattern is seasonal monoestry. The first mating period begins in January–February. Gestation is 5–6 months, and young are weaned when 1 month old. However, the presence of small- to medium-sized embryos in April–May in Sonora (Villa-R. 1966) and records of females with small embryos in January and September in Tabasco (Sánchez Hernández and Romero Almaraz 1995a) suggest that the reproductive pattern is bimodal polyestry in some parts of its geographic range. They establish maternity colonies where the bats number in the thousands. Activity. Dalquest (1953) indicated that Peters’s ghost-faced Figure 6.61. Known distribution of Peters’s ghost-faced bat (Mormoops megalophylla) in Colima.

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bats leave roosts about 2 h after sunset. Sánchez Hernández (1984) reported that they began activity at dusk and were caught until 2200 h. Individuals begin to return about 7 h after first leaving the roost (Bateman and Vaughan 1974). Other observations. Peters’s ghost-faced bats are directly beneficial to people. They often congregate in colonies as large as 500,000 individuals (Villa-R. 1966; Barbour and Davis 1969), producing large quantities of guano, which is used as fertilizer. They also consume substantial numbers of insects, some of which are detrimental to agricultural crops. It is common to find them with other mustached bats (Pteronotus) and sometimes with lesser long-nosed bats (Leptonycteris yerbabuenae). Salas-Rojas et al. (2004) assessed 3 ghost-faced bats from Colima for the presence of antirabic antibodies. One tested positive, indicating previous exposure to rabies (Lyssavirus). Sheeler-Gordon and Smith (2001) tested 2 specimens from Colima for rabies; both results were negative. In Colima, average flight speed in dense lowland forest (mangrove) was 18.6 km/h for males; females were evaluated on 2 nights, with averages being 17.3 and 20.9 km/h (Kennedy et al. 1977). In a thick upland habitat (dense, thorny vegetation), average speeds were 17.0 and 15.3 km/h for males and females, respectively (Hopkins et al. 2003).

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Measurements. Females were significantly larger than males in forearm length and maxillary-toothrow length (appendix B). Specimens examined (n = 160). 16 CNMA; 35 KU; 30 LACM; 5 LSUMZ; 1 NMNH; 61 OMNH; 11 UAEM; 1 UMMZ. Localities (fig. 6.61). 1. 1 OMNH; 14. 5 LSUMZ; 49. 1 CNMA; 52. 3 OMNH; 53. 4 OMNH; 68. 1 OMNH; 79. 1 CNMA; 88b. 1 CNMA; 91. 1 CNMA; 95. 3 OMNH; 97. 2 CNMA, 2 OMNH; 97a. 1 CNMA; 15 OMNH; 98. 1 CNMA; 98e. 1 KU; 98g. 1 CNMA; 6 OMNH; 98j. 3 OMNH; 99. 34 KU; 116b. 1 CNMA; 129. 1 NMNH; 133. 1 LACM; 142. 1 OMNH; 144f. 5 CNMA, 16 OMNH, 10 UAEM; 152. 1 OMNH; 157. 1 OMNH; 160. 1 UAEM; 164. 1 CNMA; 165b. 1 UMMZ; 166. 29 LACM; 205a. 1 OMNH; 225. 3 OMNH. Records in literature. 52, 52b, 53, 95 (Kennedy et al. 1984); 98f (Smith 1972); 98g (Kennedy et al. 1984); 99a (Davis and Carter 1962; Smith 1972); 100a (Davis and Carter 1962); 107 (Kennedy et al. 1977); 129 (Davis and Carter 1962); 144f (Hopkins et al. 2003); 154a, 157a (Kennedy et al. 1984); 163 (Sheeler-Gordon and Smith 2001); 163e (López Wilchis and Torres Flores 2005); 165b (Davis and Carter 1962); 166 (Smith 1972); 229a* (Phillips and Jones 1968).

DAVY’S NAKED-BACKED BAT

Pteronotus davyi Gray, 1838

Figure 6.62. Davy’s naked-backed bat, Pteronotus davyi.

Type locality is “Trinidad” (Gray 1838:500). Pteronotus is derived from the combination of two Greek words, pteron meaning “wing” and notos meaning “back,” referring to the wing membrane being “connected with the middle line of the back by a narrow ligament instead of arising from the sides of the body as in closely related species” (Palmer 1904:596). The specific epithet davyi honors “Dr John Davy, so well-known for his physiological studies” (Gray 1838:500).



Description. Davy’s naked-backed bat is medium-sized, with a total length of 71–85 mm and forearm length of 41–50 mm (fig. 6.62). The species has a reddish color phase and another that is dark brown. Labial and nasal skin folds are complete. The labio-nasal plate is moderately complex in structure, and the margin above each nostril is ornamented by a series of irregularly shaped, wart-like tubercles. Ears are lanceolate and funnel-shaped; the tragus is moderately complex and spatulate. Wing membranes are united mid-dorsally, giving the appearance of a hairless back and providing a possible advantage in creating an air pocket on the under surface of the wing to give added support (Goodwin 1946). The tail emerges dorsally from the uropatagium at knee level. The profile of the skull is relatively flat, and the rostrum is short and narrow, bending sharply upward; the braincase is wide. The tympanic ring is large, nearly covering the bulla, and the basioccipital region between the bullae is broad. Upper incisors are small, with a gap between incisors and canines. Lower incisors are small and trilobate. Dental formula is i 2/2, c 1/1, p 2/3, m 3/3, total 34 (Goodwin and Greenhall 1961). Distribution. Southern Baja California, southern Sonora, and Nuevo León to northwestern Peru and northern Venezuela, as well as Trinidad and southern Lesser Antilles (Simmons 2005). Elevational range sea level to 1,500 m (Villa-R. 1966). In Colima, 4–1,197 m. Conservation status. Least concern (IUCN 2014). In Mexico, no special status (SEMARNAT 2010). Habitat. Davy’s naked-backed bat was captured in mist nets set along streams and rivers under or around wild fig trees (Ficus), mango trees (Mangifera indica), and acacia bushes (Acacia). It was found in highly disturbed tropical deciduous forest, with secondary vegetation (Polaco and Muñiz-Martínez 1987). In Guerrero, the species was in a deep cave (25 mm), and claws show extreme lateral compression. The uropatagium is well developed, and the tail is more than one-half as long as the femur, with the tip of the tail emerging dorsally from the uropatagium. The calcar is well developed and bony. The skull is robust with distinct postorbital processes. The deep braincase is rounded, and the rostrum is strongly arched. Nares are somewhat tubular, opening anteriorly with a slight posterior emargination. The mastoid region conspicuously flares and is shelflike. The palate, which is concave transversely and almost flat anteroposteriorly, extends posteriorly beyond the toothrows. Bullae are relatively small, but covering about 50% of the cochlea. The skull has a well-developed sagittal crest, more so in males than females. Dental formula is i 2/1, c 1/1, p 1/2, m 3/3, total 28 (Miller 1907; Hall 1981; Hood and Jones 1984). Distribution. Sinaloa to the Guianas, southern Brazil, northern Argentina, Paraguay, Bolivia, and Peru, as well as Trinidad, Greater and Lesser Antilles, and southern Bahamas (Simmons 2005). Elevational range sea level to 250 m (Hall 1981). In Colima at 4 and 39 m. Species only moves locally (Sánchez-Hernández 1984). Conservation status. Least concern (IUCN 2014). In Mexico, no special status (SEMARNAT 2010). Habitat. Greater bulldog bats can be in many types of vegetation, from tropical deciduous forest to tropical evergreen forest. They are in coastal and disturbed areas associated with palm plantations and mangroves (Rhizophora). They congregate in groups under branches of mangroves, in hollows of trees, in abandoned houses, and in caves. They were observed and captured most frequently over ponds and slow-flowing streams, but small colonies also were associated with estuaries of large rivers and in bays (Hood and Jones 1984). In Brazil, a concrete bridge that traversed a semipermanent stream contained a nocturnal feeding roost used by more than 150 individuals (Willig 1983). Diet. The diet of greater bulldog bats is primarily fish. However, it also includes ants (Isoptera), beetles (Coleoptera), stink bugs (Pentatomidae), and cockroaches (Blattodea; Hood and Jones 1984). They eat small fish, aquatic invertebrates (e.g., shrimp), and terrestrial invertebrates (e.g., crabs; Emmons and Feer 1997). Stomach contents of three specimens in Costa Rica and Panama contained 53% fish and 47% insects (Fleming et al. 1972). In Venezuela, fish were not

Noctilio leporinus

147

et al. 1971). In Michoacán, 3 pregnant females were recorded in February (Sanchez Hernandez et al. 1985), while in Panama, 30 pregnant females were documented in February (Allen 1937). In Venezuela, there were pregnant females in June and September and lactating females in February, June, and November (Ibañez Ulargui 1981). Activity. Greater bulldog bats have been observed flying after 2100 h above rivers with a slow current (Sánchez-Hernández 1984). In Jalisco, a specimen was collected at 0500 h (López-Forment et al. 1971). In Venezuela, they were active from 1900 to 2100 h (Ibañez Ulargui 1981). In Costa Rica, an individual was recaptured 1.62 km from its original capture site (Fleming et al. 1972) and, in Venezuela, some were captured more than 2 km from the nearest trees (Ibañez Ulargui 1981). Other observations. Greater bulldog bats may roost in colonies of up to several hundred individuals (Goodwin and Greenhall 1961). In Mexico, males and females, as well as juveniles and adults, occupied a cavity made by woodpeckers 2.1 m above ground in a coconut palm (Cocos nucifera; Jones et al. 1973). In Veracruz, a colony of 16 was in a large hollow

present in stomach contents in September (when water bodies were relatively full) but made up 100% of stomach contents in November (Ibañez Ulargui 1981). Individuals seem to coordinate feeding activities and concentrate foraging activity on the periphery of water bodies (Willig 1983). Reproduction. Examined 12 adult males and 6 adult females. Male, inguinal testes, 2 (January [7×5, 7×6]). Scrotal testes, 1 (January [8×4]). Female with 1 embryo, 1 (January [5×3]). No embryo, 1 (January). The reproductive pattern may be bimodal polyestry. Breeding usually begins in November–December. The first period of births is April–June with the second in October– December (Carter 1970). Postpartum estrus can occur (Hood and Jones 1984). In Campeche, 6 young females and males were recorded in July (Jones et al. 1973). Reports of pregnant females include 3 in Chiapas in October, 1 in Panama in February, and 1 in Guatemala in February (Carter et al. 1966). In Jalisco, a female with 1 embryo (33 mm) was caught in April (López-Forment

JALISCO

Pac

Figure 6.70. Known distribution of greater bulldog bat (Noctilio leporinus) in Colima.

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an

MEXICO

COLIMA Minatitlán

MINATITLÁN 19°15'

COMALA

VILLA DE Comala ÁLVAREZ Villa de Álvarez Colima COQUIMATLÁN Coquimatlán

MANZANILLO

Cuauhtémoc

CUAUHTÉMOC

COLIMA

Manzanillo ARMERÍA

117 19°00'

Ixtlahuacán IXTLAHUACÁN 193

Armería

Tecomán TECOMÁN

Greater Bulldog Bat Noctilio leporinus

18°45'

MICHOACÁN

20 Kilometers 10 Miles N 104°45'

148

FAMILY NOCTILIONIDAE

104°30'

104°15'

104°00'

103°45'

103°30'

tree about 6–9 m above ground (Carter et al. 1966); average minimum distance between adjacent bats was about 30 cm. In Brazil, colonies were composed of 30 or fewer individuals (Willig 1983). A greater bulldog bat is known to have lived 11.5 years (Wilkinson and South 2002). In Colima in January 2002, 1 of the 5 that were caught did not have a functional right eye; it was extremely aggressive, trying to bite anyone who tried to hold it. Measurements. Males were significantly larger than females in total length, mass, greatest skull length, condylocanine length, maxillary-toothrow length, mandible length,



and mandibular-toothrow length (appendix B). There is geographic variation in size (Davis 1973); largest individuals are in northern and southern parts of the geographic range, and the smallest are in the Amazon Basin (Hood and Jones 1984). Specimens examined (n = 18). 4 CNMA; 1 NMNH; 13 OMNH. Localities (fig. 6.70). 117. 1 NMNH; 193a. 1 CNMA, 11 OMNH; 193c. 3 CNMA, 2 OMNH. Records in literature. 117 (Sánchez-Hernández et al. 2002); 193b (Kennedy et al. 1984).

Noctilio leporinus

149

Family Natalidae (Funnel-eared Bats) The family contains 3 genera and 8 species (Simmons 2005). Funnel-eared bats are insectivores that occur from northern Mexico, across Central America and the West Indies, and southward into southeastern Brazil. Dry semideciduous and secondary growth forests in tropical and semitropical lowlands and foothills are commonly occupied habitats. Colonial, funnel-eared bats typically roost in the darkest recesses of hot, moist, and deep caves and abandoned mines, but some roost in hollow trees. Day roosts may contain from just a few to hundreds of individuals. Within colonies, individuals are usually widely spaced. These are slim-bodied bats with long slender wings, tail, and legs; the legs can be longer than the head and body combined. Size ranges from 5 to 10 g, wingspan is 18–20 cm, fur is soft and long, and color of pelage usually is yellowish to cinnamon, but may be gray or brownish. Ears are large (broader than high) and shaped like three-fourths of a funnel, which probably serves to focus sounds of flying moths and other insects during foraging. Funnel-eared bats appear to be well suited for flying in habitats with a cluttered understory. Their fluttering flight is almost mothlike, and foraging flight is slow, delicate, and maneuverable (Nowak 1999; Barnett 2004; Vaughan et al. 2015). For Colima, 150 museum specimens document 1 genus and 2 species as occurring in the state, including the woolly funnel-eared bat (Natalus lanatus, 2 specimens) and Mexican greater funnel-eared bat (N. mexicanus, 148). A maternity colony of Mexican greater funnel-eared bats is highlighted in figure 6.71, with adults caring for young.

Figure 6.71. Family Natalidae. Mexican greater funnel-eared bats (Natalus mexicanus) in a maternity colony, with adults caring for young.

150

WOOLLY FUNNEL-EARED BAT

Natalus lanatus Tejedor, 2005

Figure 6.72. Woolly funnel-eared bat, Natalus lanatus.

Type locality is “6 miles SSE of Las Varas, Nayarit, Mexico” (Tejedor 2005:1110). Gray (1838) did not give reasons for choosing Natalus, which likely is derived from the Latin natalis meaning “of or relating to birth.” Goodwin (1959) believed that it might be related to the unknown origin or “birthplace” of the taxon. As pointed out by Tejedor (2011), some authors (e.g., Gómez-Laverde 1986) have speculated that Natalus has something to do with the neotenic aspect of this group of bats as in “newborn.” The specific epithet lanatus “is an anagram of the genus name Natalus, and is also Latin for woolly, in reference to the woolly texture of the pelage of the species” (Tejedor 2005:1110).



Description. The woolly funnel-eared bat is small, with a total length of 90–92 mm and forearm length of 35–39 mm (fig. 6.72). It has bicolored ventral fur; dorsal and ventral hairs are always darker at the base than at the tips. Pelage is dense, woolly, and dull, being grayish to ochraceous. The ventral surface and more than 50% of the dorsal surface of the ears are profusely covered with hair. Legs and feet are conspicuously hairy, with tufts of long hairs projecting from bases of claws. Legs are considerably shorter than the forearm. The medial margin of the ear is straight. The skull is delicately built, with an elongated and anteriorly inflated braincase. The sagittal crest is well developed, and the rostrum is shallow. Posterior margins of the maxilla are almost perpendicular to the longitudinal axis of the skull in ventral view. The basisphenoid pits are shallow and double; basisphenoid furrows are shallow but slightly deeper than the basisphenoid pits. The mandible is slender, the ascending ramus of the dentary is markedly upturned, the coronoid process is slightly lower than the condyle, relative to the alveolar plane, and the angular process is long, with an anterodorsally curved apophysis. Upper dentition is neither particularly robust nor crowded. Crowns of P2 and P3 are subequal and only slightly shorter than that of P4. Dental formula is i 2/3, c 1/1, p 3/3, m 3/3, total 38 (Tejedor 2005). Distribution. Chihuahua, Durango, Guerrero, Jalisco, Nayarit, Sinaloa, and Veracruz (Tejedor 2005), Oaxaca (Santos-Moreno et al. 2010b), Costa Rica (Tejedor 2011), and now Colima. Elevational range 50–2,000 m; most localities for which geographic coordinates are known are in middle elevations (Tejedor 2011). In Colima, at 4 m and 8 m. Conservation status. Least concern (IUCN 2014). In Mexico, no special status (SEMARNAT 2010). Habitat. The woolly funnel-eared bat inhabits a variety of climatic regimes, ranging from dry subtropical forest with marked seasonal variations in temperature and precipitation to continuously moist montane tropical forest. Distribution probably is limited more by availability of roosts than by type of vegetation. It has been collected in caves and mines. The woolly funnel-eared bat seems moderately gregarious but may roost alone; however, more than 50% of collection localities from throughout its range are represented by a single museum specimen (Tejedor 2011). In addition, it has been captured in a mist net set over a creek in a transition zone from pine-oak and tropical deciduous forests (Tejedor 2005) to moist montane tropical forest. This species may be less gregarious than other species of Natalidae. Diet. Nothing is known of the diet or activity patterns of woolly funnel-eared bats, although they probably are similar in ecology to other species of the genus in being slow-flying insectivores that forage in low vegetation (Tejedor 2005, 2011). Natalus lanatus

151

JALISCO

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Figure 6.73. Known distribution of woolly funnel-eared bat (Natalus lanatus) in Colima.

an

MEXICO

COLIMA Minatitlán

COMALA VILLA DE Comala ÁLVAREZ Villa de Álvarez Colima

MINATITLÁN 19°15'

Cuauhtémoc

CUAUHTÉMOC

COQUIMATLÁN Coquimatlán

MANZANILLO

COLIMA

97 98 Manzanillo ARMERÍA 19°00'

Ixtlahuacán IXTLAHUACÁN

Woolly Funnel-eared Bat

Armería

Tecomán TECOMÁN

Natalus lanatus

18°45'

MICHOACÁN

20 Kilometers 10 Miles N 104°45'

104°30'

Reproduction. Examined 2 adult males. No reproductive information was recorded. The reproductive pattern possibly is seasonal monoestry, as in other congeneric species. Other observations. López-Wilchis et al. (2012) noted that in Mexico the proposed diagnostic morphological characters to separate the woolly funnel-eared bat from the Mexican greater funnel-eared bat (Natalus mexicanus) are not consistent, and rarely differentiate between the two forms. They also surmised that phylogenetic analyses did not support

152

FAMILY NATALIDAE

104°15'

104°00'

103°45'

103°30'

the separation of Natalus into 2 species in Mexico. These conclusions are at variance with those of Tejedor (2011), who noted several morphological features that set the woolly funnel-eared bat apart from others in the genus. Measurements. Not able to evaluate sexual dimorphism due to having only two male specimens (appendix B). Specimens examined (n = 2). 2 OMNH. Localities (fig. 6.73). 97a. 1 OMNH; 98d. 1 OMNH. Records in literature. None.

MEXICAN GREATER FUNNEL-EARED BAT

Natalus mexicanus Miller, 1902

Figure 6.74. Mexican greater funneleared bat, Natalus mexicanus.

Type locality is “Santa Anita, Lower California, Mexico” (Miller 1902:399). Gray (1838) did not give reasons for choosing Natalus, which likely is derived from the Latin natalis meaning “of or relating to birth.” Goodwin (1959) believed that it might be related to the unknown origin or “birthplace” of the taxon. As pointed out by Tejedor (2011), some authors (e.g. Gómez-Laverde 1986) have speculated that Natalus has something to do with the neotenic aspect of this group of bats as in “newborn.” The specific epithet mexicanus is Latin, with Mexico and suffix -anus, indicating “from or pertaining to Mexico.”



Description. Mexican greater funnel-eared bats have a small body with a long tail (fig. 6.74). Total length is 87–99 mm and forearm length 35–39 mm. They have a fragile appearance. The species has four color variants—pale gray or gray in young and reddish or yellowish in adults. Pelage is silky, lax, and long (8–10 mm dorsally). When grayish (as in juveniles and young adults), it shows a distinctive patch of dark-tipped hairs posterior to the shoulders. The face and nares are simple, and eyes are small. The medial margin of the ears is slightly concave, forming an angle (250 cm/year) and the dry season, if any, is brief. Such forests are diverse floristically and structurally, with trees sometimes reaching great heights. Most trees are evergreen. Deciduous species, if present, do not all lose their leaves at the same time of year. type locality  Very specific location where a type specimen was obtained. type specimen  Specimen used in the original description of a new species or subspecies. unicuspid  Tooth with a single cusp, such as a canine tooth. urban  Of cities and larger towns; more specifically,

landscapes dominated by structures and activities of humans.



venter  Belly or underside of an organism. ventral  On lower or bottom side or surface. vertebrate  Animal (including mammals, birds, amphibians, reptiles, and fish) with a backbone. voucher material  A specimen or sample and its associated data, which document existence of an organism at a given place and time in a manner consistent with disciplinary standards. weaning  Time when a young animal stops nursing and

begins feeding on other types of food.

wing sac  Structure that functions in reproduction. Present in some species of bats in the family Emballonuridae. xerophytic scrub forest  Type of forest that may develop in an area that receives comparatively little rainfall, usually seasonally. Typically consists of shrubs and relatively short trees. young-of-the-year  An animal that was born in the most

recent breeding season and is less than a year old.

zygomatic arch  Arch of bone on the side of a mammal skull

formed by the jugal bone and a process of the squamosal bone; the “cheekbone.”

GLOSSARY

283

Figure LC.0. (above, middle, below) Abandoned building at Playa de Oro in 2000 that served as our headquarters for several of the annual expeditions to Colima.

LITERATURE CITED

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305

Figure CR.0. Cornelio SánchezHernández displays a recently captured northern yellowshouldered bat (Sturnira parvidens), as Laura Lara Ortiz (middle) and Sara GonzálezPérez (right) photograph the bat, 2007.

ILLUSTRATION CREDITS

Photographs courtesy of: Almazán-Catalán, José Alberto: 6.35, 6.57, 6.97 Altenbach, J. Scott: 6.68, 6.81, 6.91, 6.101, 6.120, 6.124, 6.126, 6.132, 6.134, 6.136, 6.138 Baker, Robert J.: 6.27 Best, Troy L.: 6.69, I.0 Daniel, Mark R.: CR.0 Geluso, Keith: 6.87 González-Pérez, Sara B.: 6.39, G.0 López Berrizbeitia, M. Fernanda: 6.83 López-González, Celia: 6.29, 6.72 Rojas-Martínez, Alberto Enríque: 6.118

Romero-Almaraz, María de Lourdes: 1.3, 1.4, 2.0, 2.9, 2.11 (left, right), 2.13 (left, right), 2.15 (left, right), 3.2 (right), 3.3, 5.0, 6.0, 6.1, 6.2, 6.8, 6.9, 6.11, 6.15, 6.17, 6.19, 6.21, 6.31, 6.33, 6.37, 6.53, 6.60, 6.62, 6.74, 6.104, 6.106, 6.116, 6.122, A.1 Ruiz-Gutiérrez, Fernando: 6.79, 6.89 Sánchez-Hernández, Cornelio: 2.8, 2.10 (left, right), 2.12 (left, right), 2.14, 6.4, 6.13, 6.23, 6.25, 6.41, 6.43, 6.45, 6.51, 6.55, 6.59, 6.64, 6.66, 6.71, 6.77, 6.85, 6.93, 6.99, 6.102, 6.108, 6.112, 6.130 Schnell, Gary D.: 1.0, 1.1 (above, below), 1.2, 3.0 (upper left, upper right, lower left, lower right), 3.1, 3.2 (left), 4.0, LC.0 (above, middle, below) Segura-Ramos, Jenifer Ruth: 6.6 Zalapa, Silvia S.: 6.47, 6.49, 6.76, 6.95, 6.110, 6.114, 6.128

307

Figure I.0. Sunset at Rancho Majagua on Pacific Ocean.

INDEX

Page numbers in italic type indicate the entry is represented in a photograph or illustration. An additional photograph of each bat is included in its section of the species accounts in chapter 6. Abies religiosa, 214 Acacia, 16, 81, 98, 112, 123, 134, 137, 190, 193; cochliacantha, 18; pennatula, 17 Acrididae, 162 adult, 277 Agavaceae, 89 Agave, 71, 73, 74, 82, 84, 85, 87, 88, 168; dasylirioides, 85; horrida, 85; inaequidens, 85; salmiana, 85 age (adult, juvenile, subadult), 26 agriculture: crops, 15, 16, 19, 136, 144, 155; land, 12, 14, 15, 18, 19; pests, 227 Aguilar C., M., 4 Akins, James, 8 albino, 97 Albizia, 81 Alcántara, Luz, 20 Alcorn, J. R., 3 alder, 76, 77, 113; Mexican, 91 alligator wood, 16 Almazán-Catalán, José Alberto, 307 Alnus, 76, 77; jorullensis, 91 Altenbach, J. Scott, 307 altricial, 277 Amaryllidaceae, 85 American Museum of Natural History, 3, 22, 30, 31, 33 Amman, Brian, 20 amole, 18 Amphipterygium adstringens, 95 anatomy, external, 41 Andropogon, 18 angelica tree, 16 Anoura geoffroyi (Geoffroy’s tailless bat), 7, 32, 37, 39, 47, 66, 70–72, 243, 249, 254, 261 ant, 59, 147, 162; flying, 164, 174, 178, 191 Antarctic, 282 anterior, 277 Anthus, 113 anticoagulant, 277 antitragus, 277

Arachis hypogaea, 98 Arachnida, 59, 211 Araneae, 223 Arbutus, 140 arch, zygomatic, 43, 283 Arctic Circle, 282 Argentinian brown bat, 7, 31, 33, 37, 39, 55, 185, 188–89, 247, 252, 257, 272 arid, 277 Armería, 9 Artibeus, 24, 36, 50, 124, 126, 130; hirsutus (hairy fruit-eating bat), 38; intermedius (intermediate fruit-eating bat), 3, 7, 24, 32, 37, 39, 40, 51, 56, 66, 109–11, 118, 245, 250, 255, 264; jamaicensis (Jamaican fruit-eating bat), 7, 32, 34, 37, 38, 39, 51, 66, 112–15, 245, 250, 255, 265; lituratus (great fruit-eating bat), 7, 32, 37, 39, 51, 66, 109, 111, 116–18, 245, 250, 255, 265; palmarum, 109; perspecillatus, 109; phaeotis, 125 ash, 196 aster, 113 Asteraceae, 113 Astrocaryum mexicanum, 61 avocado, 103 Aztec fruit-eating bat, 7, 24, 33, 37, 39, 51, 66, 123–24, 126, 245, 251, 256, 266 Aztec mastiff bat, 165 azulillo tree, 119 Baker, Robert J., 307 Balantiopteryx plicata (gray sac-winged bat), 6, 7, 32, 34, 37, 39, 46, 57, 58–60, 243, 249, 254, 260 banana, 19, 77, 90, 91, 101, 121, 123, 125, 128, 131, 198, 200, 218 banana bat, 3, 7, 32, 37, 38, 39, 48, 66, 90–92, 244, 250, 255, 263 barba de toro, 103 Barrigón, 11 basioccipital, 43, 277 basisphenoid, 43, 277

309

Bauhinia, 88; ungulata, 85 Bay: Manzanillo, 11; Santiago, 11 bee, 59 beetle, 59, 64, 138, 144, 147, 159, 161, 162, 164, 169, 173, 174, 176, 177, 183, 186, 191, 194, 197, 199, 203, 207, 210, 212, 215, 221, 223, 226, 229; ground, 169; leaf, 162; longhorned, 162; scarab, 162, 169 behavior, salting, 64 Best, Troy L., 21, 307 big brown bat, 7, 33, 37, 39, 54, 185, 190–92, 247, 252, 257, 272 big crested mastiff bat, 7, 33, 37, 39, 53, 156, 180–81, 247, 252, 257, 271 big free-tailed bat, 7, 33, 37, 39, 53, 156, 177–79, 246, 252, 257, 271 Billia hippocastanum, 103 biodiversity, 278; hotspot, 3 biome, 278 bird, 66, 183, 185 black mastiff bat, 7, 32, 37, 39, 53, 156, 166–67, 246, 252, 257, 269 black myotis, 24, 216 Blattodea, 93, 147 Boca de Pascuales, 10, 12 Bombacaceae, 85, 89 Bombax, 16, 73, 85, 88 Bonaparte, Charles Lucien, 101 Bonellia macrocarpa, 18 Boolootina, R., 4 Brazilian brown bat, 7, 23, 31, 33, 34, 35, 37, 39, 55, 185, 186–87, 271 Brazilian free-tailed bat, 7, 32, 37, 39, 52, 134, 156, 182–84, 247, 252, 257, 271 breadnut, 16, 67, 101, 106, 113 breadth: braincase, 23, 42, 254–58; mastoid, 42; rostral, 42; zygomatic, 23, 42, 254–58 breeding, seasonal, 282 broad-eared free-tailed bat, 7, 31, 33, 37, 39, 53, 156, 175–76, 184, 246, 252, 257, 270 Brosimum, 16, 101; alicastrum, 16, 67, 106, 113 bug, 169, 178, 183, 199, 221, 223; stink, 147 bugbear, 134 bulla, auditory, 43, 277 bulldog bat, 146 Buller, Audley C., 3, 34 bully, 16 Bulnesia sarmientoi, 210 Bursera, 15, 79, 90, 95, 109, 112, 190, 193 butterfly, 130, 144 cacao tree, 109, 125 Cactaceae, 85, 89 310

INDEX

cactus, 85; cardon, 218; columnar, 16, 73, 90; organ pipe, 73, 79, 82, 90, 109, 112, 113; pitahaya, 18 calcar, 41, 278 Californian leaf-nosed bat, 135, 140 Californian myotis, 7, 33, 37, 39, 52, 54, 185, 214–15, 221, 248, 253, 258, 274 Calliandra, 16, 71, 88; houstoniana, 85 Calophyllum brasiliense var. rekoi, 16 Campo Uno, 12, 17 cane, sugar, 19, 119 canine, 47, 278; lower, 42; upper, 42 canopy, 278 Capsicum, 19; annuum, 107 capsule, keratin, 45, 61 Carabidae, 169, 229 cardon, Mexican giant, 88 Carica papaya, 101, 218 Carnegiea gigantea, 87 carnivore, 66, 278 Carollia: castanea subrufa, 38; perspicillata (Seba’s shorted bat), 38, 109; subrufa (gray short-tailed bat), 6, 7, 32, 37, 38, 39, 46, 66, 101–2, 244, 250, 255, 264 Carolliinae, 7, 46, 66, 244, 250, 255 Carter, Dilford C., 216 Carter’s black myotis, 7, 24, 33, 37, 38, 39, 54, 185, 216–17, 248, 253, 258, 275 cartilage, 278 Casimiroa edulis, 113 Cassia, 16 cat, domestic, 68 cattail, 61, 161 cattle, 19, 68 caudal, 278 Cave: Aripo, 72; Tío Bartolo, 134 cave myotis, 7, 31, 33, 37, 39, 54, 185, 225–27, 248, 253, 258, 276 cazahuate, 190, 193 Cecropia, 101, 129; obtusifolia, 110, 113, 116, 122, 125, 129; peltata, 107 cedar, 103 Cedrela tonduzii, 103 Ceiba, 16, 71, 73, 81, 85, 88; aesculifolia, 85, 91; pentandra, 91, 116 Celtis reticulata, 200 cenote, 189 Centro de Investigaciones Biológicas, 22 Centurio senex (wrinkle-faced bat), 7, 33, 37, 39, 43, 46, 49, 66, 119–20, 245, 250, 255, 265 Cephalocereus, 109, 112; mezcalensis, 18 Cerambycidae, 162 Cercopidae, 174

Cerro: Chino, 11; del Otate, 11; del Zacate, 11; El Pelón, 11; Grande, 11, 12, 17, 206, 222 Chagas disease, 183 cherry, 101; capuli, 104, 123 chicozapote, 110, 113, 161 Chiroderma salvini (Salvin’s big-eyed bat), 7, 32, 37, 39, 49, 66, 121–22, 245, 250, 255, 265 Chironomidae, 197, 229 chiropatagium, 278 Choeronycteris mexicana (Mexican long-tongued bat), 7, 33, 37, 38, 39, 48, 66, 73–74, 243, 249, 254, 261 chondroalbuminoid, 278 chondromucoid, 278 Chrysomelidae, 162 cicada, 159, 177, 194, 207 Cicadellidae, 162, 174, 177 Cicadidae, 207 cingulum, 42, 278 cinnamon myotis, 7, 33, 37, 39, 54, 185, 218–19, 248, 253, 258, 275 ciruela, 95 Citrus: latifolia, 19, 106, 119; paradisi, 101; sinensis, 19 Clethra matudai, 103 Clifton, P. L., 3 climate, 12, 14 coastal, 278 cockroach, 93, 147 cocoa, 188 Cocos nucifera, 19, 61, 63, 101, 106, 109, 112, 116, 125, 128, 148, 161, 168, 175, 200, 218 Coffea arabica, 103, 109, 113, 116, 125, 129, 186 coffee, 103, 109, 113, 116, 125, 129, 186 Colección Nacional de Mamíferos, 4, 22, 31 Coleoptera, 59, 64, 96, 135, 138, 141, 144, 147, 159, 161, 164, 166, 169, 173, 177, 183, 186, 191, 194, 197, 199, 203, 207, 210, 212, 215, 221, 223, 226 Colima, city of, 3, 9, 11 colli, 9 collimaitl, 9 colonial, 278 colony, maternity, 59, 71, 72, 85, 88, 89, 96, 135, 150, 154, 177, 178, 183, 185, 192, 199, 206, 215, 229 Comala, 9 commercial value, 74, 80, 82, 91 Commissaris, Larry R., 75 Commissaris’s long-tongued bat, 7, 32, 37, 39, 49, 66, 75–76, 243, 249, 254, 261 commissure, 278 common bat, 185 common big-eared bat, 7, 32, 37, 39, 49, 66, 98–100, 244, 250, 255, 263

common mustached bat, 6, 7, 32, 34, 37, 39, 51, 133, 135, 140–42, 245, 251, 256, 267 common vampire bat, 6, 7, 32, 34, 35, 36, 37, 39, 46, 66, 67–69, 243, 249, 254, 260 congeneric, 278 conservation, 6, 18, 24, 38, 280; status, 278 conspecific, 278 constriction: interorbital, 23, 42, 254–58; postorbital, 281 Convolvulaceae, 85 Conzattia, 81 Copernicia gigas, 175 Cordia, 81; alliodora, 91 corn, 19, 121, 173, 198 cornsnake, red, 227 cortex, entorhinal, 279 Corynorhinus, 53; mexicanus (Mexican big-eared bat), 7, 31, 33, 35, 37, 38, 39, 54, 185, 205–6, 209, 247, 253, 258, 273; townsendii (Townsend’s big-eared bat), 7, 31, 33, 37, 39, 54, 185, 206, 207–9, 247, 253, 258, 274 cottonwood, 16 Coussapoa purpusii, 16 crab, 147, 155 cranium, 42, 43 Crataegus mexicana, 104, 123 crest: lambdoidal, 280; sagittal, 282 cricket, 174, 177, 178, 226 crustacean, 146, 185 Cryptocarpa procera, 95 cuachalalate, 95 Cuauhtémoc, 9, 11 cubata, 18 Cucurbitaceae, 173 cudjoewood, 18 Culicidae, 194, 199, 207, 216 culture: Aztec, 123; Toltec, 128 Cupressaceae, 200 Cupressus, 123, 196 cusps, 278; lingual, 280 Cynomops: greenhalli (Greenhall’s dog-faced bat), 158; mexicanus (Mexican dog-faced bat), 7, 33, 37, 38, 39, 53, 156, 157–58, 246, 251, 256, 268 cypress, 123, 196, 200 dactylopatagium, 41 damselfly, 199 Daniel, Mark R., 307 dark-nosed small-footed myotis, 7, 33, 37, 39, 52, 54, 185, 220–22, 248, 253, 258, 275 Davis, J. A., 3 Davy, John, 137

INDEX

311

Davy’s naked-backed bat, 3, 6, 7, 32, 34, 37, 39, 51, 133, 137–39, 140, 141, 143, 245, 251, 256, 267, 277 deciduous, 278 deforestation, 18 degradation, 278 Del Toro, 11 deme, 279 demographics, 279 Dendropanax arboreus, 16 dentary, 43, 279 dentition, 42 Dermanura, 36, 50; azteca (Aztec fruit-eating bat), 7, 24, 33, 37, 39, 51, 66, 123–24, 126, 245, 251, 256, 266; phaeotis (pygmy fruit-eating bat), 7, 24, 32, 34, 37, 39, 51, 66, 125– 27, 245, 251, 256, 266; tolteca (Toltec fruit-eating bat), 7, 24, 32, 34, 37, 39, 51, 66, 128–30, 245, 251, 256, 266 Dermaptera, 141 desert, 84, 161, 200 Desmodontinae, 7, 46, 66, 243, 249, 254 Desmodus rotundus (common vampire bat), 6, 7, 32, 34, 35, 36, 37, 39, 43, 46, 66, 67–69, 243, 249, 254, 260 devil’s-fig, giant, 110, 113, 122, 125, 129 diastema, 279 Diclidurus albus (northern ghost bat), 7, 33, 37, 39, 44, 46, 57, 61–62, 243, 249, 254, 260 diet, 36 digit, 279 dimorphism, 279; sexual, 23, 27, 243, 282 Diospyros nigra, 77, 113 Diptera, 64, 138, 141, 159, 166, 169, 173, 178, 183, 191, 194, 199, 207, 210, 212, 215, 221, 229 disc-winged bat, 38 distal, 279 diurnal, 279 diversity, 3; diet, 35 dog, domestic, 68 dorsal, 279 dorsum, 279 Dracaena, 119 dragonfly, 93, 191, 197 dragon tree, 119 ear, 41, 48; bonnet-shaped, 45 eastern small-footed myotis, 222 echolocation, 279 ecology, 279 ectoparasite, 22, 279 Elaeis oleifera, 106, 116, 128, 166 El Cóbano, 6 elevation, 10, 25, 59, 70, 104, 106, 128, 144, 194, 198 elm, Spanish, 91 312

INDEX

El Mixcoate, 4, 38 El Terrero, 17 Emballonuridae, 7, 35, 36, 44, 46, 57, 58, 243, 249, 254, 283 embryo, 26 Enchisthenes hartii (velvety fruit-eating bat), 7, 31, 33, 34, 37, 38, 39, 50, 66, 131–32, 245, 256, 266 endemic, 3, 38, 79, 90, 202, 205, 216, 279 endoparasite, 279 Enterolobium, 16, 98 entoconid, 42 Ephemeroptera, 141, 144, 203 Eptesicus, 54; brasiliensis (Brazilian brown bat), 7, 23, 31, 33, 34, 35, 37, 39, 55, 185, 186–87, 271; furinalis (Argentinian brown bat), 7, 31, 33, 37, 39, 55, 185, 188–89, 247, 252, 257, 272; fuscus (big brown bat), 7, 33, 37, 39, 54, 185, 190–92, 247, 252, 257, 272 era: Cenozoic, 11; Mesozoic, 11 Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional, 22 estrus, 279; postpartum, 281 Eucalyptus, 71 Eumops, 53; ferox (fierce bonneted bat), 7, 24, 33, 37, 39, 53, 156, 159–60, 246, 251, 256, 269; glaucinus (Wagner’s bonneted bat), 24, 160; underwoodi (Underwood’s bonneted bat), 7, 33, 37, 39, 53, 156, 161–63, 246, 252, 257, 269 Euphorbiaceae, 15 Euterpe oleracea, 93 Fabaceae, 15, 79, 85 facultative, 279 falcon, 227 Falconidae, 227 femur, 41 fertilization, 279; delayed, 278 Ficus, 4, 15, 16, 56, 63, 98, 101, 107, 109, 113, 116, 119, 121, 123, 125, 128, 129, 131, 134, 137, 161, 190, 193, 200, 218; aurea, 110, 113; palmieri, 63; pertusa, 116 fiddlewood tree, 93 Field Museum of Natural History, 3, 22, 30, 31, 33 fierce bonneted bat, 7, 24, 33, 37, 39, 53, 156, 159–60, 246, 251, 256, 269 fig: Florida strangler, 110, 113; wild, 4, 15, 16, 56, 63, 98, 101, 107, 109, 113, 116, 119, 121, 123, 125, 128, 129, 131, 134, 137, 161, 190, 193, 200, 218 fish, 36, 146, 147, 161, 185, 210, 278, 281 Flanagan, P. A., 3 Flavivirus, 155 flea, 226 fly, 64, 138, 159, 169, 173, 178, 183, 191, 194, 199, 207, 210, 212, 215, 221; caddis, 226; lacewing, 191

foramen, incisive, 43 forearm, 41 forest: cloud, 12, 70, 123, 175, 186, 197; conifer, 87, 128, 140, 166, 198; conifer-oak, 81, 103, 106, 109, 121, 128, 134, 161, 164, 171, 177, 180, 182, 188, 202, 207, 216, 218, 223; deciduous, 63, 79, 98, 106, 119, 125, 157, 166, 278; evergreen, 57, 63, 101, 131, 157, 171, 186, 188, 193, 197; gallery, 17, 101, 119, 121, 131, 186; humid tropical, 202; montane, 103, 104, 151; oak, 14, 15, 17, 87, 123, 166, 190, 193, 196, 205, 221; oak-madrone, 140; pine, 103, 112; pine-oak, 14, 15, 17, 18, 70, 73, 77, 79, 84, 93, 116, 123, 131, 151, 175, 188, 193, 196, 198, 205, 212, 214, 221, 223, 225; secondary, 67, 150, 186; semideciduous, 17, 57, 63, 79, 98, 101, 116, 131, 150; subtropical, 151, 216; subtropical moist, 159; temperate, 159, 282; tropical, 133, 137, 153, 159, 161, 216, 223; tropical deciduous, 14, 15, 16, 17, 58, 61, 67, 70, 73, 75, 77, 79, 81, 87, 90, 93, 101, 109, 112, 116, 119, 121, 128, 131, 137, 140, 143, 147, 151, 153, 164, 171, 173, 175, 177, 180, 188, 198, 202, 212, 218, 221, 225, 283; tropical dry, 77, 90, 121, 161, 173, 225; tropical evergreen, 77, 93, 106, 119, 125, 147, 153, 161, 164, 166, 173, 175, 177, 180, 186, 202, 218; tropical semideciduous, 14, 15, 16, 17, 18, 19, 58, 61, 70, 81, 82, 95, 103, 107, 109, 113, 123, 126, 130, 134, 141, 153, 157, 161, 198, 210, 214, 218; xerophytic scrub, 67, 73, 75, 79, 81, 87, 90, 98, 106, 109, 112, 123, 128, 134, 140, 161, 164, 173, 175, 177, 197, 202, 207, 212, 221, 225, 283 Formicidae, 162, 164, 174, 178, 191 formula, dental, 42, 279 fossa: anterior cranial, 277; mesopterygoid, 280 fox, gray, 68 fragmentation, 279 Fraxinus, 196 free-tailed bat, 156 fringed myotis, 7, 31, 33, 35, 37, 39, 54, 185, 223–24, 248, 253, 258, 275 froghopper, 174 frontal, 43 frugivore, 35, 36, 37, 66, 93, 99, 119, 122, 129, 131, 279 frugivore-omnivore, 36, 37, 99, 101, 104, 107, 109, 113, 116, 125 fruit, 36, 57, 71, 73, 75, 77, 80, 82, 85, 88, 146 frutescence, 125, 129 Fulgoridae, 162 funnel-eared bat, 150 garambullo, 71, 73, 85, 88 Gardner, A. L., 3, 75 gazetteer, 27, 231–42 Gecarcinus, 155 gecko, Lane’s leaf-toed, 154

Geluso, Keith, 307 genera, 279 genus, 279 Geoffroy-Saint-Hilaire, E., 70 Geoffroy’s tailless bat, 7, 32, 37, 39, 47, 66, 70–72, 243, 249, 254, 261 gestation, 279 gland: gular-thoracic scent, 160, 168, 169, 279; mammary, 280 gleaner, 96, 99, 208; foliage, 279 glory bush, 75, 77, 79 Glossophaga, 36, 48; alticola, 78; commissarisi (Commissaris’s long-tongued bat), 7, 32, 37, 39, 49, 66, 75–76, 243, 249, 254, 261; leachii (Gray’s long-tongued bat), 7, 32, 37, 39, 49, 66, 77–78, 244, 249, 254, 261; morenoi (western longtongued bat), 7, 32, 37, 38, 39, 49, 66, 79–80, 244, 249, 254, 262; soricina (Pallas’s long-tongued bat), 6, 7, 8, 32, 34, 37, 39, 49, 66, 78, 81–83, 244, 249, 254, 262 Glossophaginae, 7, 46, 47, 66, 243, 249, 254 Glyphonycteris sylvestris (tricolored big-eared bat), 7, 32, 37, 39, 49, 66, 93–94, 244, 250, 255, 263 Goldman, Edward A., 3 González B., Ciro, 3, 4 González-Pérez, Sara B., 21, 306, 307 Gould, E., 3 grapefruit, 101 grasshopper, 99, 226; short-horned, 162 grassland, 15, 106, 140, 166, 177, 207, 218, 221 Gray, E., 4 Gray, J. E., 140 gray sac-winged bat, 6, 7, 32, 34, 37, 39, 46, 57, 58–60, 243, 249, 254, 260 gray short-tailed bat, 6, 7, 32, 37, 38, 39, 46, 66, 101–2, 244, 250, 255, 264 Gray’s long-tongued bat, 7, 32, 37, 39, 49, 66, 77–78, 244, 249, 254, 261 greater bulldog bat, 7, 32, 35, 37, 39, 43, 146, 147–49, 246, 251, 256, 268 greater sac-winged bat, 7, 32, 37, 39, 46, 57, 63–65, 243, 249, 254, 260 great fruit-eating bat, 7, 32, 37, 39, 51, 66, 109, 111, 116–18, 245, 250, 255, 265 Greenhall’s dog-faced bat, 158 gregarious, 279 Gryllidae, 174, 178 guácima, 113 guano, 72, 97, 133, 136, 138, 141, 144, 154, 175, 183, 227, 279 Guarea glabra, 16 guava, 16, 99, 104, 109, 110, 134; arrayan, 113; common, 75, 79, 131 INDEX

313

guayabillo, 103 gyrus, cingulate, 278 habitat, 279 Hacienda: El Cóbano, 5; La Magdalena, 3, 90 hackberry, 200 hairy fruit-eating bat, 38 Handroanthus serratifolius, 157 hare, 147 harem, 64, 113, 186, 279 Harrison, Ed N., 90 Hart, J. H., 131 hawthorn, Mexican, 104, 123 Heliocarpus occidentalis, 113 hematophagous, 67, 279 Hemiptera, 59, 96, 141, 159, 166, 169, 178, 183, 194, 199, 203, 221, 223 hibernaculum, 208, 226, 227, 279 hibernation, 178, 209, 215, 221, 222, 227, 279 highland yellow-shouldered bat, 103 Histoplasma capsulatum, 100, 141, 183, 192 histoplasmosis, 100, 141, 183, 192 H.M.S. Starling, 103, 106 H.M.S. Sulphur, 103, 106 hoary bat, 7, 31, 33, 35, 37, 39, 55, 185, 196–97, 198, 247, 253, 257, 272 homeothermy, 279 Homoptera, 59, 141, 203 Honduran yellow-shouldered bat, 7, 24, 32, 37, 39, 50, 66, 103–5, 244, 250, 255, 264 horse, 68 hound, Molossian, 164, 166, 168 humerus, 279 hundred-year-old man, 119 Hura, 16; polyandra, 16 hydrologic region: Armería-Coahuayana, 12; Costa de Jalisco, 12 Hylocereus, 78 Hymenoptera, 59, 96, 141, 166, 178, 183, 191, 194, 199, 207, 226 hypocone, 42, 52, 279 hypoconid, 42 Ichneumonidae, 177 importance, economic, 72, 89, 91 incisor, 52, 280; lower, 42, 47, 48; upper, 42, 47, 48 indigoberry, 90 infrutescence, 104, 109, 113, 122, 280 insect, 36, 75, 80, 82, 85, 88, 91, 97, 99, 101, 107, 109, 125, 136, 141, 144, 147, 155, 166, 169, 173, 177, 191, 194, 197, 205, 207, 212, 215, 221, 227; aerial, 138, 141, 188; aquatic, 183; net-winged, 162; sucking, 59 314

INDEX

insectivore, 35, 36, 37, 57, 59, 64, 66, 73, 80, 93, 96, 133, 135, 146, 150, 151, 154, 156, 157, 159, 162, 164, 166, 169, 171, 173, 176, 177, 180, 183, 185, 186, 188, 191, 194, 197, 199, 200, 203, 205, 207, 210, 212, 215, 216, 218, 221, 223, 226, 228, 280; opportunistic aerial, 281 Instituto Politéctico Nacional, 4 intermediate fruit-eating bat, 3, 7, 24, 32, 37, 39, 40, 51, 56, 66, 109–11, 118, 245, 250, 255, 264 International Union for Conservation of Nature and Natural Resources, 24, 280 Ipomoea, 71, 73, 77, 82, 85, 88, 91, 112, 190, 193; arborescens, 85; batatas, 19; murucoides, 75, 77, 80 Islands: Revillagigedo, 9; Tres Marias, 96, 202 Isolatocereus dumortieri, 95 Isoptera, 147, 173, 177 isothermality, 25, 71, 79, 104, 129, 194, 198 IUCN Red List, 24, 38, 280, 282 jaboncillo, 103 Jamaican fruit-eating bat, 7, 32, 34, 35, 37, 38, 39, 51, 66, 112–15, 245, 250, 255, 265 jícama, 19 joso, 81 jugal, 280 Jumpiche, 11 Junta Municipal Madrid, 11 kapok, 91, 116 karyotype, 280 katydid, 93, 174, 178 keel, 280 Kennedy, Michael L., 5, 21, 30, 231 Köppen, Wladimir, 12 labial, 280 lactation, 26, 280 Lagoon: Armería, 11; Cuyutlán, 11, 15 Laguna La María, 11, 214 La Media Luna, 11, 17 La Mina, 11 lancepod, 15 Lara, Angel, 5 Lara Ortiz, Laura, 306 Lasiurus, 54; blossevillii (western red bat), 7, 33, 37, 39, 55, 185, 193–95, 247, 252, 257, 272; cinereus (hoary bat), 7, 31, 33, 35, 37, 39, 55, 185, 196–97, 198, 247, 253, 257, 258, 272; ega (southern yellow bat), 201; grayi, 198; intermedius (northern yellow bat), 7, 32, 37, 39, 55, 185, 198–99, 247, 253, 258, 273; xanthinus (western yellow bat), 7, 33, 37, 39, 55, 185, 200–201, 247, 253, 258, 273 La Yerbabuena, 6, 11

Leach, William Elford, 77 leafhopper, 162, 174 leaf-nosed bat, 66 leaf-tent, 66 Lee, M. R., 3 Leguminosae, 79 Lemaireocereus, 112 length: condylocanine, 23, 42, 249–53; ear, 23, 243–48; foot, 23, 243–48; forearm, 23, 249–53; greatest skull, 23, 42, 249–53; mandible, 23, 42, 254–58; mandibular-toothrow, 23, 42, 254–58; maxillary-toothrow, 23, 42, 249–53; tail, 23, 243–48; third-metacarpal, 23, 249–53; tibia, 23, 243– 48; total, 23, 243–48 Lepidium, 113 Lepidoptera, 61, 64, 71, 96, 135, 138, 141, 144, 159, 162, 164, 166, 169, 173, 176, 177, 183, 186, 191, 194, 197, 205, 207, 210, 211, 212, 215, 216, 221, 223, 226, 228 leporinus, 43, 44 Leptonycteris, 47; curasoae, 87; nivalis (Mexican long-nosed bat), 7, 32, 37, 38, 39, 48, 66, 84–86, 87, 244, 249, 254, 262; yerbabuenae (lesser long-nosed bat), 7, 26, 32, 37, 38, 39, 49, 66, 87–89, 136, 244, 249, 254, 262 lesser long-nosed bat, 7, 26, 32, 37, 38, 39, 49, 66, 87–89, 136, 244, 249, 254, 262 lidpod, 88 lime, 19, 106, 119 lingual, 280 lip, upper, 44, 52 little yellow bat, 7, 32, 37, 38, 39, 53, 185, 202–4, 247, 253, 258, 273 little yellow-shouldered bat, 34, 106 livestock, 16, 19, 143 lizard, 66, 185 locality, type, 283 locomotion, quadrupedal, 68, 156 locus, nuclear microsatellite, 281 Lonchocarpus, 15 long-eared myotis, 212 López Berrizbeitia, M. Fernanda, 307 López-González, Celia, 307 Lygaeidae, 169 Lysiloma, 16; candidum, 161; tergemina, 95 Lyssavirus, 68, 76, 80, 82, 89, 97, 100, 107, 110, 113, 117, 126, 130, 136, 138, 141, 144, 155, 183, 192, 195, 197, 199, 201, 204, 215, 227 Macrotus: californicus (Californian leaf-nosed bat), 135, 140; waterhousii (Waterhouse’s leaf-nosed bat), 7, 32, 37, 39, 49, 66, 95–97, 244, 250, 255, 263 Madras thorn, 116, 134, 140, 173, 190, 193, 198, 218 Magnoliaceae, 15

magnum, foramen, 43 maitl, 9 mammae, 280 Mandevilla foliosa, 75 mandible, 42, 43, 280 Mangifera indica, 19, 61, 75, 79, 103, 106, 109, 110, 112, 116, 119, 128, 134, 137, 140, 168, 193, 198, 202, 218 mango, 19, 61, 75, 79, 103, 106, 109, 110, 112, 116, 119, 128, 134, 137, 140, 168, 193, 198, 202, 218 mangrove, 12, 14, 15, 136, 138, 141, 144, 147, 175; red, 61, 109 Manilkara zapota, 106, 110, 113, 161 manjack, 81 Manzanillo, 3, 9 marsh, 280 mass, 23, 243–48 Matudaea trinervia, 103 maxilla, 42, 43, 280 maxillae, 280 maxillary, 50 mayfly, 144 mealworm, 162, 178 meatus, 280 membrane, antebrachial, 277 Meneses, Esteban, 6 mentum, 198, 280 Mephitidae, 68 mesostyle, 42 mesquite, 98, 112, 128 metacarpal, 41 metacone, 42, 280 metaconid, 42 metaloph, 280 metastyle, 42, 280 Mexican big-eared bat, 7, 31, 33, 35, 37, 38, 39, 54, 185, 205– 6, 209, 247, 253, 258, 273 Mexican dog-faced bat, 7, 33, 37, 38, 39, 53, 156, 157–58, 246, 251, 256, 268 Mexican greater funnel-eared bat, 6, 7, 8, 24, 32, 37, 39, 51, 135, 140, 150, 152, 153–55, 246, 251, 256, 268 Mexican long-nosed bat, 7, 32, 37, 38, 39, 48, 66, 84–86, 87, 244, 249, 254, 262 Mexican long-tongued bat, 7, 33, 37, 38, 39, 48, 66, 73–74, 243, 249, 254, 261 Micronycteris microtis (common big-eared bat), 7, 32, 37, 39, 49, 66, 98–100, 244, 250, 255, 263 midge, 183, 197, 229 migration, 61, 73, 81, 85, 87, 106, 119, 121, 128, 134, 140, 143, 153, 166, 171, 173, 178, 183, 193, 196, 197, 200, 202, 205, 207, 212, 223, 225, 228, 280; seasonal, 36, 73 Mimosa, 16 INDEX

315

Mine: Caleta de Campos, 153; Tiamaro, 59, 154; Tuxpan, 205, 225 mine, abandoned, 58, 73, 75, 79, 87, 90, 95, 98, 101, 109, 121, 128, 131, 133, 134, 140, 143, 150, 205, 221 molar, 280; lower, 42; upper, 42, 44 molariform, 280 Molossidae, 7, 35, 36, 45, 51, 156, 159, 172, 183, 246, 251, 256 Molossus, 36, 53; aztecus, 165; molossus (Pallas’s mastiff bat), 7, 33, 37, 39, 53, 156, 164–65, 246, 252, 257, 269; rufus (black mastiff bat), 7, 32, 37, 39, 53, 156, 166–67, 246, 252, 257, 269; sinaloae (Sinaloan mastiff bat), 7, 32, 37, 39, 53, 156, 168–70, 246, 252, 257, 270 mombin, 113; yellow, 116 monoestry, 26; asynchronous, 26, 36, 39, 68, 144, 229, 277; continuous, 68; seasonal, 26, 36, 39, 59, 62, 64, 71, 74, 85, 94, 96, 99, 135, 138, 141, 144, 152, 154, 157, 160, 162, 171, 174, 176, 178, 181, 183, 187, 191, 194, 197, 199, 201, 206, 208, 212, 215, 216, 219, 221, 224, 226, 229, 282 monotocous, 280 Moraceae, 104, 107, 125 Moreno, Alfredo, 79 Mormoopidae, 7, 35, 36, 44, 51, 133, 137, 144, 245, 251, 256 Mormoops megalophylla (Peters’s ghost-faced bat), 6, 7, 28, 32, 37, 39, 51, 133, 134–36, 140, 245, 251, 256, 267 morning glory, 71, 73, 75, 77, 80, 81, 85, 88, 91, 112 mosquito, 183, 194, 199, 207, 216 moth, 61, 64, 71, 135, 138, 144, 150, 159, 162, 164, 169, 173, 176, 177, 178, 183, 186, 191, 194, 197, 205, 207, 210, 211, 212, 215, 216, 221, 223, 226, 227, 228 Mount Orizaba, 84 municipality: Armería, 9, 240–41; Colima, 9, 239–40; Comala, 9, 232–33; Coquimatlán, 9, 17, 237–39; Cuauhtémoc, 9, 233–34; Ixtlahuacán, 9, 242; Manzanillo, 9, 234–37; Minatitlán, 9, 233; Tecomán, 9, 241–42; Villa de Álvarez, 9, 233 Muntingia, 101 Musa, 19, 77, 90, 101, 121, 123, 125, 128, 131, 198, 200, 218 muscle, occipitopollicalis, 281 Museum, British, 131 Museum of: Natural Science, 4, 22, 30, 31, 33; Southwestern Biology, 22, 30, 31; Vertebrate Zoology, 23, 30, 31; Zoology, 23, 30, 31, 33 Musonycteris harrisoni (banana bat), 3, 7, 32, 37, 38, 39, 48, 66, 90–92, 244, 250, 255, 263 mustached bat, 133 Muyil Dzibanche, 65 Myotinae, 7, 53, 185, 247, 253, 258 Myotis, 53; albescens (silver-tipped myotis), 7, 33, 37, 38, 39, 41, 54, 185, 210–11, 247, 253, 258, 274; auriculus (southwestern myotis), 7, 31, 33, 37, 39, 54, 185, 212–13, 247, 253, 258, 274; californicus (Californian myotis), 7, 33, 316

INDEX

37, 39, 52, 54, 185, 214–15, 221, 248, 253, 258, 274; carteri (Carter’s black myotis), 7, 24, 33, 37, 38, 39, 54, 185, 216– 17, 248, 253, 258, 275; ciliolabrum (western small-footed myotis), 222; evotis (long-eared myotis), 212; fortidens (cinnamon myotis), 7, 33, 37, 39, 53, 54, 185, 218–19, 248, 253, 258, 275; leibii (eastern small-footed myotis), 222; melanorhinus (dark-nosed small-footed myotis), 7, 33, 37, 39, 52, 54, 185, 220–22, 248, 253, 258, 275; nigricans (black myotis), 24, 38, 216; thysanodes (fringed myotis), 7, 31, 33, 35, 37, 39, 54, 185, 253, 223–24, 248, 258, 275; velifer (cave myotis), 7, 31, 33, 37, 39, 54, 185, 225–27, 248, 253, 258, 276; yumanensis (Yuma myotis), 7, 31, 33, 35, 37, 39, 54, 185, 228–29, 248, 253, 258, 276 Myrtillocactus, 71, 73, 85, 88 name, scientific, 282 nares, 280 naris, 280 nasal, 43, 50 Natalidae, 7, 35, 36, 46, 51, 150, 151, 246, 251, 256, 280 Natalus: lanatus (woolly funnel-eared bat), 7, 24, 31, 33, 34, 37, 39, 51, 150, 151–52, 246, 251, 256, 268; mexicanus (Mexican greater funnel-eared bat), 6, 7, 8, 24, 32, 37, 39, 51, 135, 140, 150, 152, 153–55, 246, 251, 256, 268; stramineus, 6, 24, 155 National Museum of Natural History, 3, 22, 30, 31, 33, 38 Natural History Museum of Los Angeles County, 3, 22, 30, 31, 33 Nava, J., 4 Nearctic, 3, 10, 38, 280 nectar, 36, 66, 71, 77, 79, 82, 85, 88, 90, 91, 101, 107, 113 nectarivore, 66, 73, 88, 280 Nelson, Edward W., 3 Neobuxbaumia, 90; mezcalaensis, 90, 95 neonate, 280 Neotoma micropus, 227 Neotropical, 3, 10, 38, 280 net, mist, 22, 280 Neuroptera, 96, 141, 162, 191 niche, 281 Nicotiana, 198; glauca, 161 nightshade, 101, 104, 107, 116, 125; twoleaf, 122, 129 nm (no measurement), 25 Noctilio leporinus (greater bulldog bat), 7, 32, 35, 37, 39, 43, 146, 147–49, 246, 251, 256, 268 Noctilionidae, 7, 35, 43, 146, 246, 251, 256 nocturnal, 281 northern ghost bat, 7, 33, 37, 39, 46, 57, 61–62, 243, 249, 254, 260 northern yellow bat, 7, 32, 37, 39, 55, 185, 198–99, 247, 253, 258, 273

northern yellow-shouldered bat, 7, 24, 32, 37, 39, 50, 66, 106–8, 244, 250, 255, 264, 306 nosegay, 95 noseleaf, 41, 44, 281 Nuttall, Thomas, 207 Nyctinomops, 52; aurispinosus (Peale’s free-tailed bat), 7, 33, 37, 39, 53, 156, 171–72, 174, 175, 177, 246, 252, 257, 270; femorosaccus (pocketed free-tailed bat), 7, 31, 33, 34, 37, 39, 53, 156, 173–74, 246, 252, 257, 270; laticaudatus (broad-eared free-tailed bat), 7, 31, 33, 37, 39, 53, 156, 175–76, 184, 246, 252, 257, 270; macrotis (big free-tailed bat), 7, 33, 37, 39, 53, 156, 177–79, 246, 252, 257, 271 oak, 15; Mexican red, 18; ridge, 103; savannah, 76; white, 17 Ocaña, Aurelio, 4 occipital, 43 Ocean, Pacific, 12, 15, 308 ocote, 17 Odonata, 93, 96, 141, 166, 191, 197, 199 omnivore, 66, 281 Operculina, 88 Opuntia, 128; atropes, 95 orange, 19 organ, natalid, 45, 153, 280 Ortega Huerta, Miguel A., 25 Orthoptera, 93, 96, 99, 141, 166, 177, 226 Owen, Robert D., 21 owl, 227 oyamel, 214 Pachycereus, 16, 218; pringlei, 88; weberi, 90 Pachyrhizus erosus, 19 palate, 281 palatine, 43 Pallas’s long-tongued bat, 6, 7, 8, 32, 34, 37, 39, 49, 66, 78, 81–83, 244, 249, 254, 262 Pallas’s mastiff bat, 7, 33, 37, 39, 53, 156, 164–65, 246, 252, 257, 269 palm: acai, 93; American oil, 106, 116, 128, 166; chocho, 61; coconut, 19, 61, 63, 101, 106, 109, 112, 116, 125, 128, 148, 161, 168, 175, 200, 218; date, 161; Florida thatch, 161; royal, 162 Palmillas, 11 palo: blanco, 161; Colorado, 103; María, 16; santo, 210 Pantherophis guttatus, 227 papaya, 101, 218 papelillo, 193 paracone, 42 paraconid, 42 paraloph, 281

parastyle, 42 parietal, 43 Parnell, Richard, 140 parturition, 36 patronym, 281 Patten, D. R., 3 pattern, reproductive, 36 peach, 103, 104, 225 Peale’s free-tailed bat, 7, 33, 37, 39, 53, 156, 171–72, 174, 246, 252, 257, 270 peanut, 98 Pentatomidae, 147 pepper, 101, 107; chili, 19 peppergrass, 113 period: Cretaceous, 11; Quaternary, 11; Tertiary, 11 Persea americana, 103 Peters’s ghost-faced bat, 6, 7, 28, 32, 37, 39, 51, 133, 134–36, 140, 245, 251, 256, 267 Phoenix dactylifera, 161 Phyllodactylus lanei, 154 Phyllostomidae, 7, 24, 35, 38, 41, 43, 46, 66, 243, 249, 254 Phyllostominae, 7, 43, 46, 49, 66, 244, 250, 255 phylogenetic, 281 phylopatry, 281 physiographic province, Eje Volcánico Transversal, 10 pichon, 75 Pilea, 80 pine, 71, 73, 85, 88, 113; Douglas, 17; Nicaraguan pitch, 18 pinna, 281 Pinus, 71, 73, 85, 88, 113; douglasiana, 17; maximinoi, 17; oocarpa, 18 Piper, 101; auritum, 122; hispidum, 107; lapathifolium, 107; pseudolindenii, 122 Piperaceae, 107 piscivore, 35, 36, 37, 147, 281 pitahaya, 77, 88, 90, 95 pitayo, 74 Pithecellobium dulce, 116, 134, 140, 173, 190, 193, 198, 218 plagiopatagium, 41, 281 Plains of Colima, 3, 10, 18, 34 planthopper, 162 plate: cribriform, 278; supraorbital, 282 Playa de Oro, 5, 11, 63, 231, 284 plug, vaginal, 283 plum, 119, 202 Plumeria rubra, 95 pochote, 91 pocketed free-tailed bat, 7, 31, 33, 34, 37, 39, 53, 156, 173–74, 246, 252, 257, 270 Poindexter, Cassie J., 28 pollen, 36, 71, 75, 107, 113, 125 INDEX

317

pollenivore, 35, 66, 71, 73, 82, 88, 281 pollenivore-insectivore, 36, 37, 77, 79, 81, 85, 88, 91 pollenivore-nectarivore, 75 pollination, 281 polyestrous, 281 polyestry, 26, 216; asynchronous, 78, 82, 96, 110, 113, 117, 126, 277, 278; bimodal, 26, 36, 39, 76, 80, 88, 91, 96, 102, 104, 120, 122, 123, 131, 135, 148, 162, 167, 169, 189, 203, 211, 278; continuous, 26, 36, 39, 78, 82, 107, 110, 113, 117, 126, 129, 165, 278; continuous asynchronous, 278; trimodal, 80, 211, 213 polygyny, 281; resource-defense, 113, 282 population, 281 Populus guzmanantlensis, 16 postlactating, 26, 281 potato, sweet, 19 poui tree, yellow, 157 Pouteria reticulata, 106 powder puff, 16, 71, 88 precipitation: coldest quarter, 25, 70, 75, 104, 109, 128, 194; driest month, 25, 87, 194; driest quarter, 25, 70, 104, 194; mean annual, 12, 13, 25, 59, 75, 99, 104, 128; seasonality, 25, 63, 75, 99, 113, 125, 194; warmest quarter, 25, 71, 96, 141, 144, 194; wettest month, 25, 63, 75, 96, 99, 128; wettest quarter, 25, 59, 75, 99, 104, 113, 128, 144 precocial, 281 pregnant, 26 premaxilla, 43, 281 premaxillae, 281 premolar, 281; lower, 42; upper, 42 presphenoid, 43 prickly-ash, 18 prickly pear, 95, 128 process: angular, 43; condyloid, 43; coronoid, 43; olecranon, 281; postorbital, 45, 47, 281; pterygoid, 282 procone, 42 procumbent, 47, 48, 281 promiscuous, 176, 281 Promops centralis (big crested mastiff bat), 7, 33, 37, 39, 53, 156, 180–81, 247, 252, 257, 271 propatagium, 41, 44, 277, 281 Prosopis, 98, 112, 128 protoconid, 42 protoconule, 281 province, biogeographic, 278 proximal, 281 Prunus, 119, 202; persica, 103, 104, 225; serotina, 104, 123 Pseudobombax ellipticum, 75, 77, 85, 88 Psidium, 16, 99, 104, 109, 110, 134; guajava, 75, 79, 131; sartorianum, 113 Psocoptera, 203 318

INDEX

Pteronotus, 51; davyi (Davy’s naked-backed bat), 3, 6, 7, 32, 34, 37, 39, 51, 133, 137–39, 140, 141, 143, 245, 251, 256, 267, 277; parnellii (common mustached bat), 6, 7, 32, 34, 37, 39, 51, 133, 135, 140–42, 245, 251, 256, 267; personatus (Wagner’s mustached bat), 4, 6, 7, 32, 37, 39, 51, 133, 143–45, 245, 251, 256, 267 pterygoid, 282 Pueblo Juárez, 3, 4, 38, 90 Puerta de Anzar, 11 pumpwood, 107 Punta El Carrizal, 11 Putranjiva, 119 pygmy fruit-eating bat, 7, 24, 32, 34, 37, 39, 51, 66, 125–27, 245, 251, 256, 266 Quercus, 15, 140; candicans, 17; cassipes, 17; castanea, 17, 18; elliptica, 18; lancifolia, 103; laurina, 17; magnoliifolia, 17, 18; obtusata, 17; resinosa, 17, 18; salicifolia, 18 rabies, 68, 76, 80, 82, 89, 97, 100, 107, 110, 113, 117, 126, 130, 136, 138, 141, 144, 155, 183, 192, 195, 197, 199, 201, 204, 215, 227 Rafinesque, C. S., 182, 186, 188, 190 rainforest, 57, 282; montane, 103; tropical, 283 Ranchitos ejido, 6 Rancho Majagua, 308 Randra thurberi, 90 range, 282 raspberry, Andean, 113 region: biogeographic, 3, 10, 38; climate, 14; hydrologic, 12 regurgitate, 282 relict, 282 reproduction, 282 Rhizophora, 15, 147, 175; mangle, 61, 109 Rhogeessa parvula (little yellow bat), 7, 32, 37, 38, 39, 53, 185, 202–4, 247, 253, 258, 273 ridge, presphenoid, 281 Rincón de López, 11 riparian, 17, 282 River: Armería, 10, 12, 15; Barreras, 12; Cihuatlán, 12; Coahuayana, 12; El Naranjo, 12; Marabasco, 12; Salado, 12, 81, 153 river basin: Armería, 12; Chacala-Purificación, 12; Cihuatlán, 12; Coahuayana, 12 Rodentia, 59, 66 Rojas-Martínez, Alberto Enríque, 307 Romero-Almaraz, María de Lourdes, 21, 307 roost: bachelor, 141; day, 57, 66, 112, 146, 156, 185, 278; feeding, 93, 110, 147, 279; maternity, 85, 88, 207, 224, 280; nocturnal, 281 rostrum, 45, 47

Royal Ontario Museum, 23, 30, 31 Roystonea, 162 Rubiaceae, 15, 113 Rubus glaucus, 113 Ruiz-Gutiérrez, Fernando, 8, 307 sac: interfemoral, 173; wing, 44, 46, 61, 63, 64, 283 Saccharum officinarum, 19, 119 Saccopteryx bilineata (greater sac-winged bat), 7, 32, 37, 39, 46, 57, 63–65, 243, 249, 254, 260 sac-winged bat, 57 sagittal, 282 saguaro, 87, 88 Salas-Rojas, Mónica, 20 Salix, 98 sally, 282 Salvia, 129 Salvin, Osbert, 121 Salvin’s big-eyed bat, 7, 32, 37, 39, 49, 66, 121–22, 245, 250, 255, 265 Sam Noble Oklahoma Museum of Natural History, 5, 22, 30, 31, 33 San Antonio, Colima, 11 Sánchez-Hernández, Cornelio, 21, 40, 277, 306, 307 Sánchez Vázquez, Leobardo, 20 Sanderson, I., 4 San Diego, Cerro, 11 San Gabriel, Cerro, 11 sanguivore, 35, 36, 37, 66, 67, 282 San Miguel, Cerro, 11 sapodilla, 106 sapotillo, 106 Saurauia villosa, 103 savanna, 57, 186, 188, 210 sawfly, 59 scapula, 282 scapulae, 282 Scarabaeidae, 162, 169 Schaldach, W. J., Jr., 3, 90 Schnell, Gary D., 5, 21, 231, 307 scoop, 81, 282 scorpion, 185 season: breeding, 278; dry, 12, 15, 16, 75, 79, 278, 283; wet, 12, 16, 75, 80 Seba’s short-tailed bat, 38, 109 Secretaría de Medio Ambiente y Recursos Naturales, 24 Segura-Ramos, Jenifer Ruth, 307 Senna, 15 serrate, 282 shaving-brush tree, 75, 77, 88 shrimp, 147

Sideroxylon: capiri, 77; cartilagineum, 16 Sierra: de Manantlán, 10, 12; Madre del Sur, 10, 79, 139; Madre Occidental, 139; Madre Oriental, 139; Perote, 11; Tarahumara, 58, 67, 154, 221 silk plant, 81 silver-tipped myotis, 7, 33, 37, 38, 39, 41, 54, 185, 210–11, 247, 253, 258, 274 Sinaloan mastiff bat, 7, 32, 37, 39, 53, 156, 168–70, 246, 252, 257, 270 Siphonaptera, 226 skunk, 68 slope, 11, 25, 70, 75 snake: California lyre, 174; lyre, 155 social, 282 Solanum, 101, 104, 107, 116, 125; americanum, 107; chrysotrichum, 110, 113, 122, 125, 129; diphyllum, 122, 129; ochraceoferrugineum, 107; torvum, 107 southern yellow bat, 201 southwestern myotis, 7, 31, 33, 37, 39, 54, 185, 212–13, 247, 253, 258, 274 species, 282; endangered, 38, 84, 90; near-threatened, 38; rare, 6, 22, 104, 172, 186, 210; threatened, 6, 38, 73, 84, 87, 205, 282; vulnerable, 38, 87, 90 specimen: type, 3, 24, 283; voucher, 22, 34 speed, flight, 6, 59, 68, 82, 102, 136, 138, 141, 144, 155, 156, 174, 179, 184 sphenoid, 282 Sphinga acatlensis, 18 Sphingidae, 173 spider, 223 Spondias, 113; mombin, 116 squamosal, 282 squash, 173 Stenocereus, 73, 82, 90; beneckei, 85; queretaroensis, 74; thurberi, 88, 113 Stenodermatinae, 7, 46, 49, 66, 244, 250, 255 Strigidae, 227 stripe, supraorbital, 282 Sturnira, 36, 49; hondurensis (Honduran yellow-shouldered bat), 7, 24, 32, 37, 39, 50, 66, 103–5, 244, 250, 255, 264; lilium, 24, 106, 107; lilium parvidens, 107; ludovici, 24, 103, 105; parvidens (northern yellow-shouldered bat), 7, 24, 32, 34, 37, 39, 50, 66, 106–8, 244, 250, 255, 264, 306 subterete, 282 surface, masticatory, 280 swine, 68 Tabebuia rosea, 76 Tabernaemontana tomentosa var. palmeri, 18 Tadarida brasiliensis (Brazilian free-tailed bat), 7, 32, 37, 39, 52, 134, 156, 176, 182–84, 247, 252, 257, 271 INDEX

319

tail, 41 talon, 282 tamarind, 61, 75, 79, 109 Tamarindus indica, 61, 75, 79, 109 taxa, 282 taxon, 282 Tecolapa, 11 Tecomán, 9 teeth, cheek, 42, 278 temperate, 282 temperature: annual mean, 13, 25, 71, 104, 107, 129, 194, 198; annual range, 25, 59, 70, 79, 96, 104, 107, 128, 194; maximum warmest month, 25, 71, 77, 95, 104, 129, 154, 194; mean coldest quarter, 25, 59, 71, 104, 107, 129, 194, 199; mean diurnal range, 25, 63, 70, 77, 96, 104, 128; mean driest quarter, 25, 71, 104, 107, 129, 194, 199; mean warmest quarter, 25, 59, 71, 104, 129, 194; mean wettest quarter, 25, 59, 71, 104, 107, 129, 194; minimum coldest month, 25, 59, 71, 104, 107, 129, 194, 199; seasonality, 25, 90, 101, 113, 128, 144 tempisque, 77 temporal, 43 Tenebrio molitor, 162, 178 tent: leaf, 66, 113, 128; roosting, 112, 125 Tepames, 11 terete, 282 termite, 106, 173, 177 testes: abdominal, 25, 277; inguinal, 25, 280; scrotal, 25, 282 Tettigoniidae, 174, 178 Theobroma cacao, 109, 125, 188 threatened, 282 Thrinax radiata, 161 thumb, 41, 47 Thyropteridae, 38 tibia, 41 Tibouchina, 75, 77, 80 tobacco, 161, 198 Toltec fruit-eating bat, 7, 24, 32, 34, 37, 39, 51, 66, 128–30, 245, 251, 256, 266 tongue, protractile, 281 tormentosa, 18 torpid, 160, 199, 279 torpor, 208, 225, 282 Townsend, John K., 207 Townsend’s big-eared bat, 7, 31, 33, 37, 39, 54, 185, 206, 207–9, 247, 253, 258, 274 tragus, 41, 283 Transverse Volcanic Belt, 79, 283 Trapiche, 11

320

INDEX

Trichoptera, 96, 135, 141, 203, 226 tricolored big-eared bat, 7, 32, 37, 39, 49, 66, 93–94, 244, 250, 255, 263 Trimorphodon: biscutatus, 155; vandenburghi, 174 Trophis, 104, 107, 125 tropical, 283 Tropic of: Cancer, 282; Capricorn, 282 trumpet tree, 110, 113, 116, 122, 125, 129 Trypanosoma cruzi, 183 Typha, 61, 161 Tytonidae, 227 Underwood, C. F., 161 Underwood’s bonneted bat, 7, 33, 37, 39, 53, 156, 161–63, 246, 252, 257, 269 unicuspid, 283 Universidad: Autónoma del Estado de Morelos, 22, 30, 31; Autónoma Metropolitana, Unidad Iztapalapa, 22, 30, 31; Nacional Autónoma de México, 22, 30, 31, 33 University: Auburn, 5; Louisiana State, 4, 22, 30, 31, 33; Texas Tech, 5, 23, 30, 31; Western New Mexico, 23, 30, 31 University of: Arizona, 23, 30, 31, 33, 75; California at Berkeley, 23; Kansas, 3, 22, 30, 31; Memphis, 5, 22, 30; Michigan, 23, 30, 31, 33; Nevada, Las Vegas, 22, 30, 31; New Mexico, 23; Oklahoma, 5, 22 urban, 12, 14, 15, 18, 159, 175, 177, 183, 190, 283 Urocyon cinereoargenteus, 68 uropatagium, 41, 47, 50, 52, 283 U.S.S. Peacock, 171 vegetation: halophytic, 12, 14, 15; riparian, 153, 188; secondary, 12, 14, 15, 17, 18, 19, 58, 73, 75, 79, 81, 98, 109, 119, 121, 125, 137, 140, 182, 196, 214, 218, 221 velvety fruit-eating bat, 7, 31, 33, 34, 37, 38, 39, 50, 66, 131– 32, 245, 251, 256, 266 venter, 283 ventral, 283 vertebrae, tail, 44 vertebrate, 283 Vespertilionidae, 7, 24, 31, 35, 36, 38, 46, 53, 185, 247, 252, 257, 259 Vespertilioninae, 7, 53, 185, 247, 252, 257 Villa R., Bernardo, 4, 30, 38 virus, dengue, 155 Vismia, 101 Vitex: divaricata, 93; mollis, 119 Volcán: de Fuego, 2, 10, 12, 17, 21; de Nieve, 10, 12 Volcano, Paricutín, 163, 222 voucher material, 283

Wagner’s bonneted bat, 24, 160 Wagner’s mustached bat, 4, 6, 7, 32, 37, 39, 51, 133, 143–45, 245, 251, 256, 267 wasp, 59, 191; ichneumon, 177 Waterhouse, G. R., 95 Waterhouse’s leaf-nosed bat, 7, 32, 37, 39, 49, 66, 95–97, 244, 250, 255, 263 weaning, 283 western long-tongued bat, 7, 32, 37, 38, 39, 49, 66, 79–80, 244, 249, 254, 262 western red bat, 7, 33, 37, 39, 55, 185, 193–95, 247, 252, 257, 272 western small-footed myotis, 222 western yellow bat, 7, 33, 37, 39, 55, 185, 200–201, 247, 253, 258, 273 width, rostrum, 41 willow, 98 wing, pterygoid, 48, 282



woodrat, southern plains, 227 woolly funnel-eared bat, 7, 24, 31, 33, 34, 37, 39, 51, 150, 151–52, 246, 251, 256, 268 wrinkle-faced bat, 7, 33, 37, 39, 49, 66, 119–20, 245, 250, 255, 265 young-of-the-year, 283 Yucca, 87, 214 Yuma myotis, 7, 31, 33, 35, 37, 39, 54, 185, 228–29, 248, 253, 258, 276 Zalapa, Silvia S., 307 zalate tree, 63 Zanthoxylum, 18 zapote: black, 77, 113; white, 113 Zea mays, 19, 121, 173, 198 Ziziphus mexicana, 18

INDEX

321