Table Of ContentTHE MAMMALS OF LUZON ISLAND
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THE
MAMMALS
OF LUZON
ISLAND
Biogeography
and Natural History of a
Philippine Fauna
Lawrence R. Heaney,
Danilo S. Balete, and Eric A. Rickart
Illustrated by Velizar Simeonovski
and Andria Niedzielski
JOHNS HOPKINS UNIVERSITY PRESS BALTIMORE
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Library of Congress Cataloging- in-Publication Data
Heaney, Lawrence R.
The mammals of Luzon Island : biogeography and natural history
of a Philippine fauna / Lawrence R. Heaney, Danilo S. Balete, Eric A.
Rickart ; illustrated by Velizar Simeonovski and Andria Niedzielski.
pages cm
Includes bibliographical references and index.
ISBN 978-1-4214-1837-7 (hardcover : alk. paper) — ISBN
978-1-4214-1838-4 (electronic) — ISBN 1-4214-1837-1 (hardcover :
alk. paper) — ISBN 1-4214-1838-X (electronic) 1. Mammals—Philip-
pines—Luzon. I. Balete, Danilo S., 1960– II. Rickart, Eric A. III. Title.
QL729.P5H434 2016
599.09599'1—dc23 2015008660
A catalog record for this book is available from the British Library.
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Contents
Preface vii 3: Old Endemics: Earthworm Mice 135
Acknowledgments ix
4: New Endemic Rodents 166
Part I. The Biogeography of Diversity 5: Non-Native Rodents 173
Chapter 1. The Mammals of Luzon, 1895– 2012 3 Chapter 11. A Guide to the Bats 182
Chapter 2. Climate and Habitats 15 1: Introduction to Bats 183
Chapter 3. Discovering Diversity: Topography and 2: Family Pteropodidae: Fruit Bats and Flying
Elevational Diversity Patterns 26 Foxes 186
Chapter 4. Geological History and Fossil 3: Family Emballonuridae: Sheath-Tailed Bats 208
Mammals 40 4: Family Megadermatidae: Ghost Bats and False
Chapter 5. Producing Diversity: Speciation and Vampire Bats 213
Diversifi cation 57 5: Family Hipposideridae: Roundleaf Bats 214
Chapter 6. Habitat Disturbance and Invasive 6: Family Rhinolophidae: Horseshoe Bats 224
Species 71
7: Family Vespertilionidae: Evening Bats 234
Chapter 7. Conservation of Luzon’s Mammalian
8: Family Molossidae: Free-Tailed Bats 258
Diversity 82
Chapter 8. Synthesis: Island Biogeography Theory Glossary 263
and the Mammals of Luzon 94 Literature Cited 269
Index 285
Part II. Natural History of Luzon Mammals
Chapter 9. Large Mammals 109
Chapter 10. A Guide to the Small Mammals 117
1: Order Soricomorpha: Shrews and Their
Relatives 118
2: Old Endemics: Cloud Rats 121
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Preface
Tropical islands have held great attraction for biologists We also are strongly motivated to share what we
for centuries. The reason seems to lie both in the often have learned—to share the excitement of exploration
exceptionally great concentration of unique biodiver- and discovery, and to share the new knowledge gained.
sity on such islands and in the implicit questions that Formal teaching in a classroom or speaking to a crowd
arise from that unique diversity: what species are pres- in an auditorium are fi ne ways to impart much of the
ent on the island, how and when did they arrive, why excitement and knowledge, but writing can be the best
are there so many highly distinctive species, and how way to share the information as widely as possible, and
has the unique geological history of the islands infl u- most biologists love to write. Further, we feel a strong
enced the biota that is present? compulsion to help protect the wonderful animals we
Biologists are an inherently curious lot; the bigger have studied and the places where we have studied
and more complex the questions, the happier we seem them. We fi rmly believe that protecting these animals
to be. That curiosity is what drew the authors of this and places will work to the good of people everywhere,
volume to studies of the highly diverse mammals of both those who are so fortunate as to have them in
the Philippines. The best available evidence at hand their backyard, and those who dream, as we once did,
indicates that Luzon has the greatest concentration of of green forests on steep mountains where the un-
unique species of mammals of any place in the world. known awaits.
We found the attraction of trying to understand why This book is intended to answer to all of these parts
that is the case to be wonderfully compelling. of being a biologist. It is designed to be useful as a fi eld
Biologists are also explorers, attracted to the un- guide, a classroom textbook, and a source of informa-
known and to a sense of discovery. In our case, orga- tion for anyone who needs or simply wishes to know
nizing teams of fi eld researchers and porters from local more about mammalian diversity on Luzon. It is not a
communities to trek into rugged mountains in isolated comprehensive compilation of all of what is known re-
parts of the country has an undeniable romantic ap- garding every aspect of each species, or the sum total of
peal, and we confess that it was a large part of the at- the data that we have gathered. Rather, it is a summary
traction for us. And from the fi rst time any of the three of what we know at this time about the evolutionary or-
of us captured species that no biologist had seen before, igin and ecological maintenance of mammalian diver-
the surge of adrenalin was powerful. sity on Luzon, from a biogeographical perspective. For
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VIII PREFACE
some additional details, we refer the reader to the on- not include information about the wonderful marine
line version of the Synopsis of Philippine Mammals, mammal fauna of the Philippines; aside from the Syn-
available at www. fi eldmuseum .org/ synopsis- philippine opsis, we recommend Alava et al. (1993), Dolar et al.
-m ammals/. We offer our regrets that this volume does (2006), and Tan (1995) as sources.
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Acknowledgments
This book is the result of 15 years of focused effort to phonso, Pedro Alviola, Mike Carleton, Pedro Gonzales,
document the extraordinary mammal fauna of Luzon, Karl Hutterer, Karl Koopman, Guy Musser, and Rich-
but it is also the product of far longer periods for all ard W. Thorington Jr. Silliman University provided our
of us. Heaney benefi ted greatly from early encourage- base of operations in the Philippines during much of
ment, and opportunities to study mammals and their this period and made us so welcome that it came to feel
patterns of distribution and diversity, provided by Tom like home.
McIntyre, Richard Thorington Jr., Hank Setzer, Charles The 12 years of fi eld work on which this book is pri-
Handley, and Jim Mead at the US National Museum of marily based would not have been possible without the
Natural History; their generosity with time and knowl- strong support of the Philippine Department of Envi-
edge set him securely on the road to fi eld-o riented ronment and Natural Resources (DENR), especially the
studies of mammalian diversity, including an appre- Protected Areas and Wildlife Bureau (recently renamed
ciation for the mammals of Southeast Asia. Rickart’s the Biodiversity Management Bureau). We offer our
initial interest in mammals stems from his early asso- most sincere thanks to all of the current and former
ciation with Charles Remington and his son Eric, who staff, especially to Angel C. Alcala, T. Mundita Lim, Cora
revealed the joys of fi eld exploration, the value of close Catibog-S inha, Luz Gonzales, Samuel P eñafi el, Wil-
observation, and the powerful attraction of natural his- fredo Pollisco, Josie DeLeon, Carlo Custodio, Marlynn
tory and science as lifelong pursuits. Heaney and Rick- Mendoza, and Anson Tagtag. A great many other staff
art both studied under Robert Hoffmann, whose gentle members of the DENR in Regions 1, 2, 3, 4, 5, and the
demeanor, love for learning, and deep interest in bio- Cordillera Autonomous Region gave us their help and
geography and mammals has provided a lifetime of in- support; they are far too numerous to name, but to all
spiration. Balete was inspired by many teachers and col- of them we offer our gratitude. We thank the staff of the
leagues, especially Lipke Holthuis and Boris Sket, and National Commission for Indigenous People (NCIP) for
benefi ted immeasurably from the generosity of Anton their support in obtaining the necessary permission for
Cornelius Jacobus Burgers. our studies.
Our early studies of biogeography and mammals in We have also benefi ted from a long, close, and
the Philippines, beginning in the 1980s, were encour- highly productive association with the National Mu-
aged, supported, and inspired by a great many people: seum of the Philippines, especially Jeremy Barns, Anna
foremost among them were Angel C. Alcala, Pablo Al- Labrador, Angel Bautista, Julie Barcelona, Arvin Dies-
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