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>psis of the Mammalian Fauna
Philippine Islands

rencc R. Heaney

.

ilo S. Balete

William L. R. r

.ouella Dolar

Perry S. Ong

Alcala

Eric A. Rickart

res T. L. Dans

Bias R. '

Gonzales

Ruth C. B. Ltzurrum

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FTELDIANA

Zoology

NEW SERIES, NO. 88

A Synopsis of the Mammalian Fauna
of the Philippine Islands

il Lawrence R. Heaney

IDepartment of Zoology

Field Museum of Natural History
\ Roosevelt Road at Lake Shore Drive

Chicago, Illinois 60605-2496 USA

Danilo S. Balete Myrissa V. Lepiten

M. I < nulla Dolar William L. R. Oliver

Angel C. Alcala Perry S. Ong

Andres T. L. Dans Eric A. Rickart

Pedro C. Gonzales Bias R. Tabaranza, Jr.

Nina R. Ingle Ruth C. B. Utzurrum

Note: Remaining authors ' affiliations and current addresses may he found on page Hi.

Accepted August 12, 1997
Published June 30, 1998
Publication 1493

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 1998 Field Museum of Natural History

ISSN 0015-0754

PRINTED IN THE UNITED STATES OF AMERICA

Contributors

Lawrence R. Heaney

Department of Zoology
Field Museum of Natural History
Roosevelt Road at Lake Shore Drive
Chicago, Illinois 60605-2496 USA

Danilo S. Balete

Department of Zoology
Field Museum of Natural History
Roosevelt Road at Lake Shore Drive
Chicago, Illinois 60605-2496 USA

Current address:
Haribon Foundation
9A Malingap Street
Teachers Village
Diliman 1 1 00
Quezon City, Philippines

M. Louella Dolar

Department of Biology
Silliman University
6200 Dumaguete City
Negros Oriental, Philippines

Current address:

Scripps Institute of Oceanography

9500 Gilman Drive

University of California at San Diego

San Diego, California 92093-0204 USA

Angel C. Alcala

Silliman University Office
636 T. M. Kalaw Street
Ermita, Manila, Philippines

Current address:

Commission on Higher Education

DAP Building

Pasig City, Metro Manila, Philippines

Andres T. L. Dans

Institute of Biological Sciences
College of Arts and Sciences
University of the Philippines at Los Banos
College, Laguna, Philippines

Pedro C. Gonzales

Department of Zoology
Philippine National Museum
Rizal Park
Manila, Philippines

Nina R. Ingle

Department of Natural Resources

and New York Cooperative Fish and Wildlife

Research Unit
Cornell University
Ithaca, New York 14853 USA

Myrissa V. Lepiten

Center for Tropical Conserx'ation Studies
Silliman University
6200 Dumaguete City
Negros Oriental, Philippines

William L. R. Oliver

Flora and Fauna International
Great Eastern House
Tenison Road
Cambridge CB1 2DT
United Kingdom

Perry S. Ong

Institute of Biology

University of the Philippines

Diliman

Quezon City, Philippines

Eric A. Rickart

Utah Museum of Natural History

University of Utah

Salt Lake City, Utah 84112 USA

Bias R. Tabaranza, Jr.

Department of Biology

Iligan Institute of Technology

Mindanao State University

Iligan City, Lanao del Norte, Philippines

Current address:

Birdlife Philippines-Haribon

9A Malingap Street

Teachers Village

Diliman 1100

Quezon City, Philippines

Ruth C. B. Utzurrum

Center for Tropical Conserx'ation Studies
Silliman University
6200 Dumaguete City
Negros Oriental, Philippines

Current address:

Department of Marine and Wildlife Resources

P.O. Box 3730

Pago Pago, American Samoa 96799

in

Table of Contents

Abstract 1

Introduction 1

Content of the Synopsis 2

Distributional Patterns of Philippine

Mammals 2

Fossil Mammals 4

Procedures Used in Compiling This Syn-
opsis 4

Common Name 4

Distribution 5

Habitat 6

Status 7

Official/Legal Status 7

A Note on Authorship 8

Insectivora 8

Erinaceidae — Hedgehogs and Gymnures .... 8

Soricidae — Shrews 9

Scandenti a 11

Tupaiidae — Tree Shrews 11

Dermoptera 12

Cynocephalidae — Flying Lemurs 12

Chiroptera 12

Pteropodidae — Fruit Bats 12

Emballonuridae — Sheath-tailed Bats 20

Megadermatidae — False Vampire and

Ghost Bats 21

Rhinolophidae — Horseshoe and Roundleaf

Bats 21

Vespertilionidae — Vesper and Evening

Bats 25

Molossidae — Free-tailed Bats 29

Primates 30

Loridae — Lorises and Coucangs 30

Tarsiidae — Tarsiers 30

Cercopithecidae — Monkeys 30

Pholidota 31

Manidae — Pangolins 31

Rodentia 31

Sciuridae — Squirrels 31

Muridae — Mice and Rats 33

Hystricidae — Porcupines 44

Carnivora 44

Felidae — Cats 44

Mustelidae — Weasels, Otters, and

Badgers 45

Herpestidae — Mongooses 45

Viverridae — Civets 46

Artiodactyla 46

Suidae — Pigs 46

Tragulidae — Mouse-deer 47

Cervidae — Deer 47

Bovidae — Cattle 48

Cetacea 48

Balaenopteridae — Rorquals 48

Delphinidae — Ocean Dolphins 49

Kogiidae — Dwarf and Pygmy Sperm

Whales 51

Physeteridae — Sperm Whales 52

Ziphiidae — Beaked Whales 52

Phocoenidae — Porpoises 52

Sirenia 53

Dugongidae — Dugongs and Sea Cows 53

Discussion and Conclusion 53

Acknowledgments 54

Literature Cited 55

List of Illustrations

1 . Map of Philippine Islands 3

2. Map of Luzon showing boundaries of
provinces 5

3. Map of Mindanao showing boundaries of
provinces 6

4. Heads of representative mammals from
Mindanao 10

5. Heads of representative Philippine bats ... 13

6. Heads of representative murid rodents

from Luzon 33

7. The leopard cat, Prionailurus bengalensis .. 45

A Synopsis of the Mammalian Fauna of the
Philippine Islands

Lawrence R. Heaney and Collaborators

Abstract

The mammalian fauna of the Philippine Islands is now known to include 201 species: 22
are marine mammals, 172 are native terrestrial mammals, and 7 are introduced, non-native
species that occur in the wild. With 1 1 1 (64%) of the terrestrial native species as endemics,
the archipelago has one of the highest per-area levels of endemism in the world, on the basis
of both absolute numbers and percentage. Since the last checklist was published, in 1987, 16
new species have been discovered — currently one of the highest rates of discovery in the world
(Morell, 1996); 14 of the species listed here are not yet formally described. For each species,
we provide the citation for the original description (except when the species has yet to be
formally described), the English common name, the documented distribution, a summary of
habitat data, and an assessment of conservation status. At least 52 native species are threatened
(many seriously endangered) as a result of destruction of forest habitats, over-hunting, and
destruction of cave and marine ecosystems. Further basic research and implementation of ef-
fective habitat protection programs, especially for the forests, are essential to protect this re-
markably diverse and endangered fauna.

Introduction

The mammalian fauna of the Philippine Islands
is remarkably diverse and species rich. As docu-
mented in this study, the terrestrial fauna is now
known to include 172 native species (plus 7 in-
troduced species, most of which are widespread
in the country), which is one of the highest levels
of diversity on a per-area basis in the world. An
additional 22 species of marine mammals have
also been recorded. Moreover, most of the species
are found nowhere else: of the 172 terrestrial spe-
cies, 1 1 1 (64%) are endemic. As noted by Cole
et al. (1994) in their review of global patterns of
mammalian diversity, only Madagascar has a
higher percentage of unique species. However,
only about 80 of the 100 mammal species on
Madagascar are endemic, and Madagascar has
twice the land area of the Philippines (Heaney,
1993). Although no comprehensive analysis has
yet been done, we believe that the Philippines
have the highest level of endemic mammalian

species richness of any country on a per-unit-area
basis.

These species include some of the most spec-
tacular radiations of mammals. The murid rodent
fauna of Luzon has long been considered to be
one of the most remarkable assemblages of mam-
mals (Thomas, 1898). To name only a few, there
exist giant arboreal animals with long, lush coats
of shiny black fur (Oliver et al., 1993); excep-
tionally long-snouted, hopping rodents that feed
principally on earthworms in mossy forest (Rick-
art et al., 1991); tiny brown mice that dig through
leaf litter in their search for insects (Rickart et al.,
1991); and beautiful mice with large dark eyes,
pure white bellies, and auburn heads and backs
that scamper along forest floor and through tree-
tops (Balete & Heaney, in press). It is not sur-
prising that these creatures have generated great
interest over the century since biologists first
learned of their existence from the native peoples
of the Central Cordillera of Luzon.

The Philippines also are exceptional in a sec-
ond sense. As we report here, recent field work

FIELDIANA: ZOOLOGY, N.S., NO. 88, JUNE 30, 1998, PP. 1-61

has revealed the presence of 16 new species of
mammals in the last 10 years, since the last check-
list was published (Heaney et al., 1987). This rate
of discovery is matched in only a few other coun-
tries (Brazil and Peru), and those are more than
ten times as large as the Philippines (Morell,
1996; Wilson & Reeder, 1993). Because all of
these newly discovered species are endemic, the
estimate of the rate of endemism for the Philip-
pines has risen and likely will continue to rise.

Finally, and most important, the mammalian
fauna of the Philippines is exceptional in a third
sense. The most recent International Union for the
Conservation of Nature (IUCN) Red Data List
(Baillie & Groombridge, 1996) includes 49 threat-
ened mammals from the Philippines, which is sev-
enth in total number for any country but first
when the sizes of the countries are considered.
The recognition that the Philippines have the most
extensively endangered mammalian fauna in the
world is supported by the even more recent Phil-
ippine Red Data List (Wildlife Conservation So-
ciety of the Philippines, 1997), which lists 52 spe-
cies as endangered in the country. One of the pri-
mary purposes of this synopsis is to provide con-
sistent information on the conservation status of
all mammalian species in the country, with an in-
dication of the limitations of current information,
so as to serve as a guide both to current conser-
vation efforts and to research that will provide
even better information in the future.

Content of the Synopsis

We have compiled an updated checklist of all
mammalian species known to occur in the Phil-
ippines, including 14 as yet undescribed species.
We include, for the first time, a complete listing
of the marine mammals known or likely to occur
in the Philippines, based on a large amount of
recent field work by M. L. Dolar and her col-
leagues. This checklist contains updated and more
detailed information on the distribution of each
species than has been available previously, and
for the first time we have added brief synopses of
the habitat associations of every species.

Under species entries, we have cited relevant

publications that have appeared since the last
checklist (Heaney et al., 1987), along with critical
earlier references. For additional earlier publica-
tions, readers should consult that publication
(Heaney et al., 1987) and the annotated bibliog-
raphy of Balete et al. (1992).

Distributional Patterns of Philippine Mam-
mals — In describing the distributions of species,
we often refer to the zoogeographic regions of the
country; these are shown in Figures 1-3 (from
Heaney, 1986). The map in Figure 1 is based on
the extent of the islands in the Philippines during
the late Pleistocene period, when the development
of immense continental glaciers had, in effect, re-
moved water from the world's oceans, so that sea
level was 120 m lower than it is today (Heaney,
1991a). Each of these Pleistocene islands defines
the limit of a distinct fauna (Alcala, 1976; Dick-
erson, 1928; Everett, 1889; Heaney, 1985b, 1986).
The three largest faunal regions thus defined are
the Luzon Faunal Region (including Luzon, Ca-
tanduanes, Marinduque, Polillo, and several small
islands), the Mindanao Faunal Region (including
Mindanao, Basilan, Bohol, Leyte, Samar, and ad-
jacent small islands), and the Palawan Faunal Re-
gion (including Palawan, Balabac, Busuanga, Cu-
lion, Cuyo, and adjacent small islands). Some oth-
er islands coalesced into islands larger than those
of today (e.g., there was a single island from
Tawi-tawi to Jolo, and an island including Negros,
Panay, Cebu, and Masbate), whereas other islands
remained isolated (e.g., Camiguin, Sibuyan, Si-
quijor, and Mindoro).

The recognition of these faunal regions is an
important aid in understanding the present distri-
butions of Philippine mammals. However, our
knowledge of distribution is far from complete.
Many small islands have never been surveyed,
and many moderately large islands are poorly
known. New distributional records are discovered
by virtually every regional survey, particularly
among the more poorly known taxa (especially
bats and rodents). Further field work is essential
to determine the distributions and habitat require-
ments of many species, to document the faunas of
the smaller island groups, and to carry out the
alpha-taxonomic studies that are necessary to de-

Fig. 1. Map of the Philippine Islands, showing the locations of most of the islands mentioned in the text. Areas
within the -120 m bathymetric line are indicated; these show the limits of islands during the most recent "ice age"
and correspond to the limits of Philippine faunal regions.

FIELDIANA: ZOOLOGY

n Batanes
^ Islands

Babuyan

126

200km

Less than 120 meters
current water depth

Pacific
Ocean

South

China

Sea

y GREATER
>/ LUZON

j )* — Catanduanes

GREATER^
PALAWAN ^sk>

Maripipi
Visayan ^ _ mm

GREATER
SULU 122

Celebes
Sea

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

termine species limits and to discover new spe-
cies.

The distribution of marine mammals in this
synopsis is based on surveys, published records
of occurrence, strandings, fishery by-catch, and
skeletons found on beaches. Not all of the marine
habitats have been surveyed, and thus the absence
of records of a given species from any given area
may result from the lack of surveys rather than
real absence. Most surveys have focused on the
waters of the Visayan region such as the eastern
Sulu Sea, Tanon Strait, Bohol Sea, Panay Gulf,
and parts of the Visayan Sea (see Fig. 1 for their
locations). Other areas recently surveyed include
the southern Sulu Sea, Davao Gulf, and parts of
the Celebes Sea in the southern Philippines. No
thorough survey has been done in northern Phil-
ippine waters, although reports of cetacean sight-
ings and strandings from this region have been
included in this checklist.

Fossil Mammals

As noted in the last checklist of Philippine
mammals (Heaney et al., 1987), very little is
known about the fossil mammals of the Philip-
pines. All known species are large-bodied (ele-
phants, rhinoceros, deer, pig, and buffalo; Fox &
Peralta, 1974; Groves, 1984, 1985); their large
size contributes to the likelihood of both preser-
vation and discovery. Smaller species are cur-
rently entirely unknown, although they undoubt-
edly were present. The discovery of new material
is certain to add an exciting new dimension to our
understanding of the history of the fauna, and
may provide clues to the long-term interactions
between humans and the native fauna of the coun-
try as well as provide the means to test many of
the current models of speciation, colonization, and
extinction for Philippine mammals (e.g., Heaney,
1986, 1991b).

Procedures Used in Compiling This Synopsis

As with our prior checklist (Heaney et al.,
1987), this paper is intended as a working list of
the mammals of the Philippines. We have not pro-
vided full taxonomic synonymies; earlier papers
(Alcasid, 1970; Taylor, 1934) should be consulted
for this information. We have used Wilson and
Reeder (1993) as our starting point for species

names, taxonomic authorities, and higher catego-
ries.

We have not included subspecies designations
in this synopsis, for two reasons. First, the use of
subspecies names implies detailed knowledge of
geographic variation, and such information is
rarely available for Philippine mammals. Virtually
all subspecies names now in use need to be crit-
ically reevaluated. Second, we believe that use of
subspecies would distract the reader from more
crucial issues of general distribution and conser-
vation status.

Common Name — Because common names of-
ten assist in increasing public interest in animals,
we have chosen in this paper to provide English
common names for Philippine mammals. In one
respect, this has been a relatively easy matter: be-
cause English is a foreign language, there were
virtually no English common names in use until
about 100 years ago. English names have thus
been coined almost entirely by biologists. With a
few exceptions, we have chosen not to include
names in the many languages of the Philippines
because many local names are in use, with dif-
ferent names often used for the same species in
different linguistic areas, and sometimes a single
name is in use for different species in different
linguistic areas. Most of the few that we have in-
cluded are coming into use as English names
(e.g., kagwang and tamaraw). A complete com-
pilation of native names would be a worthwhile
and instructive effort but is beyond the scope of
this paper.

In choosing English common names to include
here, we have retained most names that are well
established; these are for species that are wide-
spread in Asia, marine mammals, or large, con-
spicuous species. Some small species that occur
outside the country have established English com-
mon names, but often several have been used. In
such cases we have retained the names where we
felt they were suitable, but in some instances we
have coined new names that are more informative
and more consistent with the biology of the ani-
mals. For the many species for which no English
names are in use, we have coined names. In doing
so, we have tried to make the names informative
about the ecology, distribution, or appearance of
the species. In some instances this has simply in-
volved translating the Latin species name where
that is suitably descriptive. We have avoided long
names and patronyms, and we have attempted to
make names interesting to the public, for whom
these names are primarily intended. Where two

FIELDIANA: ZOOLOGY

Pangasman

Less than 120 meters
current water depth

Luzon Provinces

Zam- '.Pampanga,'.'
lbal * s Vj /Bolaean i

Sataan^XM'anila'

o p^r- *>

Fig. 2. Map of Luzon showing the boundaries of provinces in 1995.

roughly equivalent common names are available,
we list both, with our preferred name listed first.
Distribution — The distribution of many spe-
cies of Philippine mammals is poorly known. This
is true primarily because little effort has been
made to survey many islands, and even less effort
has been made in the mountainous areas to which
many species are restricted (e.g., Mindanao: San-
guila & Tabaranza, 1979; Tabaranza, 1979). This
problem is being addressed by ourselves and oth-
ers, and is critically important to forming a de-
tailed and accurate picture of mammalian distri-
bution in this geographically and ecologically
complex archipelago. In this synopsis, we restrict
our statement of distribution to a summary of the
islands on which each species has been found,
with the exception of Luzon and Mindanao; on
those two large islands we report each province
in which a species has been found (provincial
boundaries are mapped in Figs. 2 and 3). The
provinces that are shown are those that existed
during the late 1980s and early 1990s when we
conducted our studies; readers should be aware
that provinces in the Philippines are split as the

human population grows, and so their boundaries
are unstable. For example. South Cotabato Prov-
ince was split in two in late 1995. with the north-
ern portion retaining the same name and the
southern being named Sarangani Province.

The term "faunal region" refers to the centers
of endemism that are defined by the extent of
Pleistocene islands shown in Figures 1-3. The six
largest faunal regions are Greater Luzon, Greater
Mindanao, Greater Palawan, Greater Negros-Pa-
nay, Greater Mindoro, and Greater Sulu, but oth-
ers (e.g., Sibuyan and Camiguin) often have en-
demic species as well. General distributions of
species can be envisioned by referring to the ac-
companying maps (Figs. 1-3).

In addition to the problems posed by insuffi-
cient surveys, many speciemens in collections in
the United States and the Philippines, including
some that formed the basis for past publications,
were mistakenly identified. This is not surprising,
given the difficulties inherent in identifying many
species, the lack of opportunity for comparison
with type specimens, and the poor quality of some
of early descriptions. However, it is a serious

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

Mindanao Provinces

) o

Less than 120 meters
current water depth

Fig. 3. Map of Mindanao showing the boundaries of provinces in 1995.

problem that requires very careful work in the fu-
ture and skeptical review of earlier publications.
We have taken a conservative approach to this
problem in this synopsis: with few exceptions, we
accept identifications only of those specimens that
have been examined and identified unequivocally
by the authors. Virtually all have been seen by the
first author. The few exceptions involve species
that are very poorly known, for which the pub-
lished record forms virtually the only information
available, or for some equivalent crucial reason.
For each Philippine island or province listed in a
species entry, we have indicated a single museum
containing at least one specimen whose identifi-
cation we have verified. Additional records from
the same islands or provinces often are available
in additional museums, but we cite only one. Mu-
seums are indicated by the following abbrevia-
tions:

amnh — American Museum of Natural History,
New York.

anm — Australian National Museum, Sydney.

bmnh — Natural History Museum, London.

cuvc — Cornell University Vertebrate Collec-
tions, Ithaca.

dmnh — Delaware Museum of Natural History,
Greenville.

fmnh — Field Museum of Natural History, Chi-
cago.

fsm — Florida State Museum of Natural History,
Gainesville.

mcz — Museum of Comparative Zoology, Har-
vard University, Cambridge.

mmnh — Bell Museum of Natural History, Uni-
versity of Minnesota, St. Paul.

msu-iit — Mindanao State University-Iligan In-
stitute of Technology, Iligan City.

pawb — Protected Areas and Wildlife Bureau,
Quezon City.

pnm — Philippine National Museum, Manila.

rom — Royal Ontario Museum, Toronto.

smf — Senckenberg Museum, Frankfurt.

su — Museum of Natural History, Silliman Uni-
versity, Dumaguete.

suml — Marine Laboratory, Silliman University,
Dumaguete.

uimnh — University of Illinois Museum of Nat-
ural History, Urbana.

ummz — University of Michigan Museum of Zo-
ology, Ann Arbor.

upd — Museum of Birds and Mammals, Univer-
sity of the Philippines at Diliman, Quezon City.

uplb — Museum of Natural History, University
of the Philippines at Los Banos, College.

usnm — United States National Museum of Nat-
ural History, Washington, D.C.

Habitat — This section of each species account
is a brief summary of the primary habitats utilized

FIELDIANA: ZOOLOGY

by the species. In each case, we indicate the types
of forest in which the species has been docu-
mented to occur, the elevational range, and the
relative abundance in each of the major habitats,
to the extent that these are known. Whenever pos-
sible, published sources are cited for the infor-
mation, but we also relied heavily on our personal
observations where these significantly supple-
mented published sources. These comments are
based entirely on specimens we believe to be cor-
rectly identified.

The principal natural terrestrial habitats in the
Philippines occur along elevational gradients,
with lowland forest, montane forest, and mossy
forest as the primary elements. The elevational
range of the habitats varies among mountains
based on peak elevation, topography, and annual
rainfall. The habitats occur lower on low moun-
tains, near the coast, and in areas with high rain-
fall and higher on mountains that have high peaks,
are far from the shore, and are relatively dry.
Lowland forest is dominated by species of the
dipterocarp family; these trees often have large
buttresses and often reach 40 m or more in height.
Lowland forest ranges from sea level to 700 m on
small, low-lying islands and to 1500 m on large,
mountainous islands. Montane forest is not dom-
inated by a single family of trees. It has trees that
typically lack buttresses and reach 12 m to 30 m
in height, and it occurs from 700 m to 1000 m on
small, low-lying islands and from 1500 m to
about 2300 m on large, mountainous islands.
Mossy forest is characterized by short, gnarly
trees (usually 3-8 m), by having moss covering
most tree trunks and branches, and by having
most ground surfaces covered by moss over a
thick layer of humic soil. Conifers with low height
but large girth (up to 2 m dbh) are found in some
mossy forests. "Heath" vegetation, dominated by
woody shrubs, often occurs on exposed ridges
within mossy forest. Mossy forest occurs from as
low as 1000 m (rarely 800 m) on small islands
(at the peaks) and from 2100 m and above on the
highest mountains. For a given mountain range,
the elevation of these types of forest may overlap,
and there is always a fairly gradual transition be-
tween them. Note that for some species of mam-
mals we know only the elevational range, not the
actual habitat in which the species occurred.

We use "primary forest" to refer to forest that
has not been disturbed significantly by major hu-
man activities such as logging. "Secondary for-
ests" have been disturbed, often heavily, usually
by logging or burning; they range from well-re-

generated second growth of natural forests, to
planted forests of exotic species, to scrubby areas
with scattered trees. We use "agricultural areas"
to refer to a range of habitats, from coconut palm
plantations to mixed pasture, fields, and orchards,
and varying densities of human habitation.

Status — This section provides a brief state-
ment of the conservation status of each species
based on published information and our personal
observations. For each species, we indicate our
best estimate of the general stability of the spe-
cies's population(s), abundance, and degree of
vulnerability to known threats (especially habitat
destruction and hunting). Where the information
is available, we indicate how a species's present
status differs from that in the past (Gonzales &
Alcala, 1969; Rabor, 1966, 1968).

We use the following terms to describe relative
abundance, in order of increasing abundance: ex-
tinct, rare, uncommon, moderately common, com-
mon, and abundant. "Endemic" refers to a spe-
cies that is restricted to a defined area; thus, a
species might be said to be endemic to Camiguin
Island, which means that it is found nowhere else.
More generally, such a species could also be said
to be endemic to the Philippines. "Native" spe-
cies occur naturally in the Philippines; all endemic
species are native, but not all native species are
endemic. For example, the palm civet (Paradox-
urus hermaphroditus) is native to the Philippines,
but it is also present in much of South and South-
east Asia. "Non-native" (also called "exotic")
species are those that do not occur naturally in the
Philippines; rather, they have been introduced into
the Philippines by humans, often accidentally,
such as the rats and mice that arrived on ships.
"Commensal" species are those that are often
(but not always) closely associated with human
settlements. These commensal species often live
in human houses and other buildings; except for
a few species of bats, all such species in the Phil-
ippines are non-native. All marine mammals listed
here are native to the Philippines, but none are
endemic.

Officiai./Lhgai- Status — Many of the species
listed herein are included in various official list-
ings of species of conservation concern and/or
protected status under national or international
agreement. These include CITES (Convention on
International Trade in Endangered Species),
IUCN (International Union for the Conservation
of Nature), and U.S. ESA (United States Endan-
gered Species Act). We include reference to all of
these listings in this synopsis, current through De-

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

cember 1996; as more is learned about the con-
servation status of the mammals of the Philip-
pines, many more species will be added to these
lists.

Users of this synopsis should be aware that the
intent and coverage of each of these lists varies
greatly. Coverage by the U.S. ESA is especially
limited with respect to Philippine species, because
the U.S. ESA deals primarily (but not exclusively)
with species in the United States and its territo-
ries. Species are listed by CITES only if it is be-
lieved that these species are threatened by inter-
national trade. It should be noted that CITES lists
some species that are moderately common in the
wild but are believed to be highly vulnerable if
trade were to be unrestricted, such as African el-
ephants. On the other hand, CITES does not list
many species that are seriously endangered, be-
cause they are not believed to be threatened by
international trade. Even species that are threat-
ened by local trade are not included by CITES if
there is no international trade. Additionally,
CITES lists species that are common if there is
reason to believe that an endangered species
might be misidentified as that species (thus in-
creasing the likelihood of trade); for example, all
species of Pteropus are listed by CITES because
several species are endangered as a result of in-
ternational trade and most species are quite sim-
ilar in appearance. In short, CITES listing is really
not a threatened species status category at all, but
a regulatory mechanism for international trade.
U.S. ESA listing are also potentially misleading,
but for different reasons. Both CITES and U.S.
ESA listings are often inconsistent with IUCN
listings, and they can be positively misleading if
misunderstood.

The IUCN Red List (Baillie & Groombridge,
1996) is the most comprehensive and consistent
of all of these listings; it is intended to cover all
parts of the world and all groups of mammals (as
well as other taxa) equally. However, it too is lim-
ited. In the past, attention was often focused on
large and attractive ("charismatic") animals, be-
cause these were often the only species for which
adequate information was available; only in the
1996 list are smaller species given equal attention.
However, even the 1996 list has limitations. First,
data are often lacking on the conservation status
(and basic ecology) of small mammals in the trop-
ics. Second, incorrect information in past lists has
sometimes been carried over (such as the listing
of Podogymnura truei, which is probably not
threatened, as discussed below). Third, the IUCN

does not include species that have been discov-
ered but have not yet been formally described. In
the case of the Philippines, where 14 species are
currently in the process of being described (as list-
ed below), this constitutes a major portion of the
fauna.

Finally, it should be noted that the Philippines
is one of the first countries in the tropics to have
its own Red Data Book (Wildlife Conservation
Society of the Philippines, 1997). The mammal
section in that book was taken largely from an
earlier draft of this paper and represents an effort
to present a comprehensive statement of current
knowledge about the conservation status of the
various species. However, as noted in that volume
and in this paper, information on many species is
very incomplete, and neither publication should
be taken as more than one step in the process of
acquiring the needed data.

A Note on Authorship — The bulk of this pa-
per was written by the first two authors, with the
exception of the sections on marine mammals,
which were written principally by M. L. Dolar.
Other authors reviewed the manuscript, made
available unpublished notes, and/or made avail-
able specimens in collections they care for. All
authors reviewed the manuscript and worked to
develop a consensus on the many issues that are
summarized here.

Insectivora

Erinaceidae — Hedgehogs and Gymnures

The only two species of the genus Podogym-
nura (Fig. 4 A) are confined to the Philippines (Po-
duschka & Poduschka, 1985).

Podogymnura aureospinula Heaney and
Morgan, 1982. Proc. Biol. Soc. Washing-
ton, 95:14.

Common Name — Dinagat gymnure, Dinagat
wood shrew.

Distribution — Endemic to the Philippines; re-
stricted to Dinagat Island (dmnh).

Habitat — Occurs in forest (Heaney & Rabor,
1982; Tabaranza, unpubl. data).

Status — Moderately common in primary and
secondary forest on Dinagat, but geographically
restricted to an island where habitat destruction

FIELDIANA: ZOOLOGY

has been extensive (Heaney & Morgan, 1982;
Heaney & Utzurrum, 1991; Tabaranza, unpubl.
data). IUCN: Endangered.

Status — Abundant and stable in continental
Asia but unknown in the Batanes group.

Podogymnura truei Mearns, 1905. Proc.
U.S. Natl. Mus., 28:437.

Crocidura beatus Miller,
U.S. Natl. Mus., 38:392.

1910. Proc.

Common Name — Mindanao gymnure, Minda-
nao wood shrew.

Distribution — Endemic to the Philippines; re-
stricted to Mindanao Island (Bukidnon [fmnh],
Davao del Norte [fmnh], and Davao del Sur
[fmnh] provinces).

Habitat — Widespread on Mindanao in primary
forest above 1300 m (Musser & Heaney, 1992),
common in montane forest, ca. 1600 m to 2000
m, and abundant in mossy forest, 2000 m to 2900
m (Hoogstraal, 1951; Poduschka & Poduschka,
1985; Sanborn, 1952; Rabor, 1986; Heaney et al.,
unpubl. data).

Status — Because of its occurrence in high-el-
evation forest of low stature that has limited com-
mercial value and its abundance, current popula-
tions of this species are stable. For these reasons,
we disagree with the IUCN listing, and we rec-
ommend that it be listed as nonthreatened (Hea-
ney et al., 1997). IUCN: Vulnerable (but we rec-
ommend delisting).

Common Name — Common Mindanao shrew.

Distribution — Endemic to the Philippines;
widespread in Mindanao Faunal Region and on
Camiguin. Recorded from Biliran (usnm), Bohol
(usnm), Camiguin (fmnh), Leyte (usnm), Maripipi
(ummz), and Mindanao (Agusan [dmnh], Bukid-
non Ifmnh], South Cotabato [uimnh], Misamis
Occidental [fmnh], and Zamboanga del Sur
[fmnh] provinces) (Heaney & Ruedi, 1994).

Habitat — Common in primary forest, especial-
ly at higher elevations; uncommon in secondary
forest, and absent outside of forest (Heaney et al.,
1989; Heaney & Tabaranza, 1995; Rickart et al.,
1993).

Status — Widespread and locally abundant, sta-
ble. IUCN: Vulnerable (but we recommend de-
listing).

Comment — Includes C. parvacauda (Heaney &
Ruedi, 1994).

Soricidae — Shrews

Systematics, genetic variation, and phylogenet-
ic variation in Philippine shrews were studied by
Heaney and Ruedi (1994). Six of the eight species
are endemic, one is widespread elsewhere in Asia,
and one is a non-native species that often lives in
and near houses and occasionally in primary for-
est. A representative species is shown in Figure
4F.

Crocidura attenuate! Milne-Edwards,
1872. Rech. Hist. Nat. Mamm, p. 263.

Common Name — Indochinese shrew.

Distribution — Widespread on Asian continent
and many associated shallow-water islands. In the
Philippines, known only from Batan Island, Ba-
tanes Province (usnm; Heaney & Ruedi, 1994).

Habitat — Common and widespread in Asia; its
ecology unknown in the Batanes group of islands.

Crocidura grandis Miller, 1910. Proc.
U.S. Natl. Mus., 38:393.

Common Name — Greater Mindanao shrew.

Distribution — Endemic to the Philippines; re-
stricted to Mindanao Island (Misamis Occidental
Province). Known only from a single specimen
(usnm) taken in 1906 at 6100 ft. on Mt. Malin-
dang (Heaney & Ruedi, 1994).

Habitat — Unknown; probably confined to pri-
mary forest.

Status — Unknown; no attempt has been made
to assess the status in the area of the type locality.
The lack of specimens from the highest peaks on
Mindanao, Mt. Apo (Sanborn, 1952) and Mt. Ki-
tanglad (Heaney et al.. unpubl. data), may indicate
that the species is confined to the Zamboanga
Peninsula and perhaps to Mt. Malindang, the
highest point on the peninsula. Deforestation has
been extensive throughout this region. Probably
vulnerable; surveys are needed. IUCN: Endan-
gered.

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

<*2

B

Fig. 4. Heads of representative mammals from Mindanao (not to the same scale). A, Podogymnura truei (Family
Erinaceidae). B, Urogale everetti (Family Tupaiidae). C, Sundasciurus philippinensis (Family Sciuridae). D, Batomys
salomonseni (Family Muridae). E, Cynocephalus volans (Family Cynocephalidae). F, Crocidura beatus (Family
Soricidae).

10

FIELDIANA: ZOOLOGY

Crocidura grayi Dobson, 1890. Ann.
Mag. Nat. Hist., ser. 6. 6:494.

Common Name — Luzon shrew.

Distribution — Philippines only; widespread in
Luzon and Mindoro faunal regions. Recorded
from Catanduanes (usnm), Luzon (Aurora [upd],
Benguet [usnm], Camarines Sur [fmnh], Laguna
[bmnh], Mountain [bmnh], and Rizal [amnh] prov-
inces), and Mindoro (mmnh). Also reported from
Abra Province, Luzon (Lawrence, 1939).

Habitat — Common in primary lowland, mon-
tane, and mossy forest from 250 m to 2400 m,
uncommon in secondary forest (Heaney et al.,
1991, in press; Rickart et al., 1991).

Status — Stable; widespread and common.
IUCN: Vulnerable (but we recommend delisting).

Comment — Includes C. halconus as a junior
synonym (Heaney & Ruedi, 1994).

habitat destruction (Heaney & Utzurrum, 1991).
IUCN: Critically endangered.

Crocidura palawanensis Taylor, 1934.
Monogr. Bur. Sci. Manila, 30:88.

Common name — Palawan shrew.

Distribution — Endemic to the Palawan Fau-
nal Region. Records are from Palawan (fmnh) and
Balabac (usnm) islands.

Habitat — The holoytpe was taken "in deep
forest near the sea" (Taylor, 1934). Also taken in
shrubby second growth at ca. 400 m (Hoogstraal,
1951). The closely related C. fuliginosa and C.
baluensis occur in forest over a broad elevational
range (Payne et al., 1985).

Status — Unknown; geographically restricted
to an area undergoing rapid deforestation. Surveys
are needed. IUCN: Vulnerable.

Crocidura mindorus Miller, 1910. Proc.
U.S. Natl. Mus., 38:392.

Common Name — Mindoro shrew.

Distribution — Philippines only; endemic to
Mindoro (usnm) and Sibuyan (fmnh) islands
(Heaney & Ruedi, 1994).

Habitat — On Mindoro, taken only in high-el-
evation primary forest; on Sibuyan, uncommon in
forest from 325 m to 1325 m.

Status — Known on Mindoro only from two
specimens taken in 1906 at 6300 ft. elevation on
Mt. Halcon, and on Sibuyan from six specimens
taken in 1989 and 1992 (Goodman & Ingle, 1993;
Goodman & Heaney, unpubl. data). Mindoro and
Sibuyan have undergone extensive deforestation.
Probably vulnerable; surveys are needed. IUCN:
Endangered.

Crocidura negrina Rabor, 1952. Chicago
Acad. Sci. Nat. Hist. Misc., 96:6.

Common Name — Negros shrew.

Distribution — Philippines only; endemic to
Negros Island (fmnh).

Habitat — Known only from six specimens
taken in primary lowland and montane forest from
ca. 500 m to 1450 m elevation in southern Negros
(ummz) (Heaney & Ruedi, 1994; Heaney & Ut-
zurrum, 1991; Rabor, 1986).

Status — Rare because of restricted range and

Suncus murinus (Linnaeus, 1766). Syst.
Nat., 12th ed., 1:174.

Common Name — Asian house shrew.

Distribution — Asia and Indo-Australia;
throughout the Philippines. Specimens from Ca-
miguin (dmnh), Leyte (usnm), Luzon (Batangas
[usnm], Cagayan [uplb], Camarines Sur [upd], II-
ocos Norte [usnm], Isabela [usnm], Laguna
[usnm], La Union [usnm], Pampanga [usnm], Pan-
gasinan [usnm], Rizal [usnm], and Zambales
[usnm] provinces), Marinduque (pnm), Mindanao
(Misamis Oriental [uplb] province), Negros
(usnm), Panay (usnm), and Siquijor (su). Also re-
ported from Caluya Island (Alcala & Alviola,
1970).

Habitat — Abundant in urban and agricultural
areas (Rabor, 1977, 1986), often in disturbed for-
est, occasionally in primary forest; found from sea
level to 1650 m on Negros (Heaney et al., 1989;
Heaney et al, in press; Rickart et al., 1993).

Status — Non-native. Abundant and stable.

Comment — Includes Crocidura edwardsiana
(Heaney & Ruedi, 1994).

Scandentia

Tupaiidae — Tree Shrews

This family contains only two Philippine spe-
cies; a representative is shown in Figure 4B.

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

11

Tupaia palawanensis Thomas, 1894.
Ann. Mag. Nat. Hist., ser. 6, 9:251.

Common Name — Palawan tree shrew.

Distribution — Endemic to Palawan Faunal Re-
gion; recorded from Balabac (usnm), Busuanga
(fmnh), Culion (fmnh), Cuyo (fmnh), and Pala-
wan (fmnh).

Habitat — Common in secondary and primary
forest, coconut groves, bamboo thickets, and ba-
nana plantations (Dans, 1993; Hoogstraal, 1951;
Sanborn, 1952) at "low to medium elevation"
(Rabor, 1986).

Status — Common and stable. IUCN: Vulner-
able (but we recommend delisting).

Comment — Included in T. glis by Corbet and
Hill (1992) but not by Wilson (1993).

Urogale everetti (Thomas, 1892). Ann.
Mag. Nat. Hist., ser. 6, 9:250.

Common Name — Mindanao tree shrew.

Distribution — Philippines only; endemic to the
Mindanao Faunal Region. Recorded from Dinagat
(dmnh), Mindanao (Bukidnon [fmnh], Davao del
Sur [fmnh], Misamis Occidental [fmnh], Misamis
Oriental [fmnh], South Cotabato [amnh], Surigao
del Norte [dmnh], and Zamboanga del Sur [usnm]
provinces), and Siargao (dmnh).

Habitat — Scarce to common in primary forest
from 750 m to 2250 m (Musser & Heaney, 1992;
Sanborn, 1952; Heaney et al., unpubl. data).

Status — Widespread and locally common in
forest on Dinagat and Mindanao, but dependent
on lowland forest that has been largely destroyed.
IUCN: Vulnerable. CITES: Appendix II.

Dermoptera

Cynocephalidae — Flying Lemurs

The order Dermoptera includes a single family,
and that family has only two species, one of
which occurs in the Philippines and the other in
Malaysia and western Indonesia. The appearance
of the head is shown in Figure 4E.

Distribution — Philippines only; endemic to the
Mindanao Faunal Region. Recorded from Basilan
(amnh), Biliran (usnm), Bohol (fmnh), Dinagat
(usnm), Leyte (usnm), Mindanao (Agusan del
Norte [uplb], Bukidnon [fmnh], Davao del Norte
[fmnh], Davao del Sur [fmnh], Lanao del Norte
[usnm], Lanao del Sur [dmnh], Misamis Occiden-
tal [fmnh], Misamis Oriental [dmnh], South Cot-
abato [fmnh], Surigao del Sur [uplb], Zamboanga
del Norte [fmnh], and Zamboanga del Sur [dmnh]
provinces), Samar (fmnh), and Siargao (dmnh).
Also reported from Maripipi (Rickart et al., 1993).

Habitat — Common in primary and secondary
forest, and in mixed forest and orchard, from sea
level to ca. 500 m elevation on small islands, up
to ca. 1 100 m on Mindanao (Rabor, 1986; Rickart
et al., 1993; Wischusen et al., 1992, 1994; Wis-
chusen & Richmond, 1989; Heaney et al., unpubl.
data).

Status — Widespread and common, popula-
tions are stable. Widespread destruction of low-
land forest makes them somewhat vulnerable, but
their ability to persist in disturbed forest makes
them more resilient than many species. IUCN:
Vulnerable (although current data do not support
this listing).

Chiroptera

Pteropodidae — Fruit Bats

The number of fruit bats known from the Phil-
ippines has increased from 23 to 25, with the ad-
dition of as yet undescribed species of Haplo-
nycteris from Sibuyan and Pteropus from Min-
doro and the new record of Pteropus dasymallus
from the Babuyan Islands, along with the inclu-
sion of Acerodon lucifer as a synonym of Acer-
odon jubatus (as discussed below). Distribution
patterns were analyzed by Heaney (1991b), Hea-
ney and Rickart (1990), and Koopman (1989),
feeding ecology by Utzurrum (1995), and chro-
mosomes by Rickart et al. (1989a). The appear-
ance of a representative species is shown in Fig-
ure 5A.

Cynocephalus volans (Linnaeus, 1758).
Syst. Nat., 10th ed., 1:30.

Common Name — Kagwang, Philippine flying
lemur.

Acerodon jubatus (Eschscholtz, 1831).
Zool. Atlas, part 4:1.

Common Name — Golden-crowned flying fox.
Distribution — Endemic to the Philippines;

12

FIELDIANA: ZOOLOGY

s^oiyjw*

..J'S/''*

Fig. 5. Heads of representative Philippine bats (not to the same scale). A, Rousettus amplexicaudatus (Family
Pteropodidae). B, Megaderma spasma (Family Megadermatidae). C, Rhinolophus sp. (Family Rhinolophidae). D,
Taphozous melanopogon (Family Emballonuridae). E, Chaerophon plicata (Family Molossidae). F, Miniopterus
schreibersii (Family Vespertillionidae). From Ingle and Heaney (1992).

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

13

widespread with the exception of the Palawan
Faunal Region and the Batanes and Babuyan
groups. Recorded from Basilan (mcz), Bongao
(amnh), Cabo (usnm), Bohol (fmnh), Dinagat
(dmnh), Jolo (fmnh), Leyte (fmnh), Luzon (Abra
[mcz], Isabela [amnh], Quezon [usnm], Rizal
[amnh], and Tarlac [usnm] provinces), Marindu-
que (upd), Maripipi (usnm), Mindanao (Davao del
Norte [fmnh], Davao del Sur [fmnh], Lanao del
Norte [dmnh], Lanao del Sur [dmnh], Maguindan-
ao [fmnh], Misamis Oriental [dmnh], South Cot-
abato [usnm], Sultan Kudarat [amnh], Surigao del
Sur [dmnh], Zamboanga del Norte [fmnh], and
Zamboanga del Sur [amnh] provinces), Mindoro
(usnm), Negros (fmnh), Panay (fmnh), Sibutu
(dmnh), and Siquijor (fmnh).

Habitat — Primary and secondary lowland for-
est up to 1 100 m. Some roosts reported from man-
grove and on small islands (Hoogstraal, 1951; Ra-
bor, 1986). Common reports of 100,000 individ-
uals in a colony from the late 1800s and early
1900s contrast with recent observations of maxi-
mum colony size of 5,000 and usually far fewer
(Heaney & Heideman, 1987; Heaney & Utzur-
rum, 1991; Lawrence, 1939; Mickleburgh et al.,
1992; Mudar & Allen, 1986; Rickart et al., 1993;
Taylor, 1934; Utzurrum, 1992).

Status — Severely declining as a result of hab-
itat destruction and heavy hunting. IUCN: Endan-
gered. CITES: Appendix II.

Comment — Includes the Panay golden-crowned
flying fox (Acerodon lucifer Elliot, 1896). Ex-
amination of all known extant specimens of A.
lucifer has shown that there are no morphological
differences that distinguish the two supposed spe-
cies (Ingle & Heaney, 1992; Heaney, unpubl.
data). Specimens from Panay show the greatest
resemblance to those from Negros. The Panay
population, which formerly occurred in large col-
onies, was presumed extinct (Elliot, 1896; Heaney
& Heideman, 1987; Mickleburgh et al., 1992; Ut-
zurrum, 1992), but recent sightings on Boracay
(Maro & Ingle, unpubl. data) may represent this
species. Possibly extinct on Siquijor (Lepiten,
1995).

Acerodon leucotis (Sanborn, 1950). Proc.
Biol. Soc. Washington, 63:189.

Common Name — Palawan flying fox.
Distribution — Philippines only; endemic to the
Palawan Faunal Region. Recorded from Balabac

(usnm), Busuanga (fmnh), and Palawan (fmnh) is-
lands.

Habitat — Uncertain, but probably moderately
common in primary and secondary forest (Hoogs-
traal, 1951).

Status — No current information. IUCN: Vul-
nerable. CITES: Appendix II.

Comment — Formerly placed in the genus Pter-
opus (Musser et al., 1982).

Alionycteris paucidentata Kock, 1969.
Senckenberg. Biol., 50:322.

Common Name — Mindanao pygmy fruit bat.

Distribution — Philippines only; endemic to
Mindanao Island (Bukidnon Province [fmnh]).

Habitat — Known only from primary forest on
Mt. Kitanglad, Bukidnon Province. Absent in
lowland forest. Uncommon in montane forest
from 1600 m to 1900 m, common in mossy forest
at 2250 m (and probably above; Heaney et al.,
unpubl. data).

Status — Common in appropriate high-eleva-
tion habitat; geographically restricted. IUCN: Vul-
nerable.

Cynopterus brachyotis (Miiller, 1838).
Tijdschr. Nat. Gesch. Physiol., 5:146.

Common Name — Common short-nosed fruit
bat.

Distribution — Southeast Asia; throughout the
Philippines. Specimens from Balabac (fmnh),
Barit (fmnh), Basilan (ummz), Batan (usnm),
Batu-bato (dmnh), Biliran (usnm), Bohol (usnm),
Bongao (dmnh), Busuanga (fmnh), Calauit
(ummz), Camiguin (fmnh), Catanduanes (fmnh),
Cebu (fmnh), Culion (fmnh), Cuyo (fmnh), Dal-
upiri (fmnh), Dinagat (usnm), Fuga (fmnh), Gui-
maras (ummz), Leyte (usnm), Luzon (Albay
[fmnh], Aurora [upd], Bulacan [usnm], Cagayan
[ummz], Camarines Sur [fmnh], Isabela [fmnh],
Laguna [usnm], Quezon [ummz], Rizal [upd], Sor-
sogon [fmnh], Tarlac [mcz, usnm], and Zambales
[usnm] provinces), Marinduque (pnm), Maripipi
(usnm), Marsec (usnm), Masbate (su), Mindanao
(Bukidnon [fmnh], Davao del Norte [fmnh], Da-
vao del Sur [fmnh], Davao Oriental [uplb], Lanao
del Norte [dmnh], Misamis Oriental [dmnh],
South Cotabato [amnh], Zamboanga del Norte
[uplb], and Zamboanga del Sur [dmnh] prov-
inces), Mindoro (fmnh), Negros (fmnh), Palawan

14

FIELDIANA: ZOOLOGY

(fmnh), Panabulon (ummz), Panay (su), Polillo
(fmnh), Sabtang (usnm), Sanga-sanga (dmnh),
Siargao (dmnh), Sibutu (dmnh), Sibuyan (fmnh),
Simunol (dmnh), Siquijor (fmnh), and Tablas (su).
Also reported from Caluya, Sibay, Semirara, Bor-
acay, and Carabao islands (Alcala & Alviola,
1970).

Habitat — Ranges from sea level to at least
1250 m. Abundant in agricultural areas, common
in secondary forest, usually uncommon or absent
in primary forest (Heaney et al., 1989, 1991, in
press; Heideman & Heaney, 1989; Ingle, 1992;
Lepiten, 1995; Rickart et al., 1993).

Status — Abundant and geographically wide-
spread; stable.

Comment — Kitchener and Maharadatunkamsi
(1991) and Schmitt et al. (1995) considered pop-
ulations of this species from the Philippines and
Sulawesi to represent a separate species, C. lu-
zoniensis, but Corbet and Hill (1992) and Koop-
man (1993) place C. luzoniensis as a subspecies
of C. br achy otis; further study is needed.

Dobsonia chapmani Rabor, 1952. Chi-
cago Acad. Sci. Nat. Hist. Misc., 96:2.

Common Name — Negros naked-backed fruit
bat.

Distribution — Philippines only; endemic to
Negros (fmnh) and Cebu (dmnh) islands.

Habitat — Formerly common in lowland forest
from sea level to 800 m in southern Negros Is-
land, where it roosted exclusively in caves (Hea-
ney & Heideman, 1987; Rabor, 1986; Utzurrum,
1992).

Status — Believed to have become extinct in
1970s as a result of the combination of forest de-
struction, disturbance by guano miners, and hunt-
ing (Heaney & Heideman, 1987; Utzurrum,
1992).

Comment — Considered to be conspecific with
D. exoleta by Corbet and Hill (1992) but not by
Koopman (1993).

Dyacopterus spadiceus (Thomas, 1890).
Ann. Mag. Nat. Hist., ser. 6, 5:235.

Common Name — Dayak fruit bat.

Distribution — Sumatra and the Malay Penin-
sula; in the Philippines, known from one speci-
men from Luzon (Abra province, smf) and one

from Mindanao (Misamis Oriental province,
dmnh) (Kock, 1969a; Utzurrum, 1992).

Habitat — Unknown in the Philippines. On
Borneo, believed to roost in hollow trees and
caves (Payne et al., 1985).

Status — Unknown.

Eonycteris robusta Miller, 1913. Proc.
Biol. Soc. Washington, 26:73.

Common Name — Philippine nectar bat, Philip-
pine dawn bat.

Distribution — Endemic to the Philippines;
widespread but absent from Palawan Faunal Re-
gion and the Batanes/Babuyan group of islands.
Recorded from Biliran (ummz), Catanduanes
(usnm), Leyte (dmnh), Lubang (mcz), Luzon
(Abra [mcz], Cavite [ummz], Mountain [mcz], and
Rizal [usnm] provinces), Marinduque (pnm), Mar-
ipipi (ummz), Mindanao (Bukidnon [fmnh], Lanao
del Norte [uplb], Maguindanao [fmnh], Misamis
Oriental [dmnh], and Zamboanga del Sur [dmnh]
provinces), Negros (usnm), and Siargao (dmnh).

Habitat — Until the 1960s, commonly taken in
caves adjacent to forest and commonly netted in
and adjacent to primary forest from sea level to
1 100 m, often in areas with mixed forest and
clearings, but never in primarily agricultural ar-
eas.

Status — Captured rarely in the 1980s and
1990s and may now be quite rare (Heaney et al.,
1991; Mickleburgh et al., 1992; Utzurrum, 1992).

Comment — Usually included as a subspecies of
E. major (Corbet & Hill, 1992; Koopman, 1993),
but we believe that it is distinct (Heaney, unpubl.
data).

Eonycteris spelaea (Dobson, 1871). Proc.
Asiat. Soc. Bengal, p. 105, 106.

Common Name — Common nectar bat, common
dawn bat.

Distribution — India to Timor; throughout the
Philippines except the Batanes/Babuyan region.
Recorded from Biliran (ummz), Bohol (usnm), Ca-
tanduanes (usnm), Cebu (uplb), Leyte (usnm), Lu-
zon (Abra [mcz], Cavite [mcz), Ilocos Norte
[usnm], Laguna [uplb], Pampanga [usnm], Rizal
[mcz], and Sorsogon [fmnh] provinces), Marin-
duque (mcz), Maripipi (ummz), Masbate (su),
Mindanao (Agusan del Norte [su], Davao del Sur
[fmnh], Davao Oriental [dmnh], Lanao del Norte

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

15

[dmnh], Misamis Oriental [dmnh], South Cotabato
[dmnh], Surigao del Norte [su], and Zamboanga
del Sur [dmnh] provinces), Mindoro (mcz), Ne-
gros (fmnh), Palawan (fmnh), Polillo (fmnh), San-
ga-sanga (dmnh), Satang (uplb), Siargao (dmnh),
Sibuyan (fmnh), Siquijor (su), and Tablas (su).
Also reported from Carabao Island (Alcala & Al-
viola, 1970).

Habitat — Common in agricultural areas from
sea level to 1100 m, often at high densities (Hea-
ney et al., 1989, in press; Heideman & Heaney,
1989; Lepiten, 1995; Rickart et al., 1993). Known
to roost only in caves, where they form colonies
of up to thousands of individuals, and where they
are vulnerable to hunting pressure (Rickart et al.,
1993; Utzurrum, 1992).

Status — Stable and common in agricultural ar-
eas but heavily hunted and vulnerable.

Haplonycteris fischeri Lawrence, 1939.
Bull. Mus. Comp. Zool., 86:33.

Common Name — Philippine pygmy fruit bat.

Distribution — Endemic to the Philippines;
widespread excluding the Camiguin, Palawan,
Batanes/Babuyan, and Sibuyan Faunal Regions.
Recorded from Biliran (usnm), Bohol (usnm), Ca-
tanduanes (fmnh), Dinagat (usnm), Leyte (usnm),
Luzon (Aurora [upd], Cagayan [ummz], Camari-
nes Sur [usnm], Isabela [amnh], Laguna [uplb],
Quezon [ummz], and Tarlac [usnm] provinces),
Marinduque (upd), Mindanao (Agusan del Norte
[ummz], Bukidnon [fmnh], Davao del Norte
[uplb], Davao del Sur [uplb], Davao Oriental
[uplb], Misamis Occidental [uplb], Misamis Ori-
ental [uplb], South Cotabato [uplb], Surigao del
Norte [uplb], Surigao del Sur [uplb], Zamboanga
del Norte [uplb], and Zamboanga del Sur [dmnh]
provinces), Mindoro (fmnh), Negros (fmnh), and
Panay (su). The record from Palawan reported by
Kock (1969b) probably is erroneous.

Habitat — One of the most common fruit bats
in primary forest, especially at middle elevations.
It is rare in secondary forest and absent in entirely
agricultural areas. Within forest, it occurs from
about 150 m to 2250 m; abundance usually in-
creases with elevation up to about 1200 m to 1500
m and then declines (Heaney et al., 1989, 1991,
in press; Heideman & Heaney, 1989; Rickart et
al., 1993).

Status — Currently stable, but it has declined in
recent decades as a result of habitat destruction

by logging. IUCN: Vulnerable (but we recom-
mend delisting).

Comment — Genetic variation was studied by
Peterson and Heaney (1993) and reproductive bi-
ology by Heideman (1989).

Haplonycteris sp. A

Common Name — Sibuyan pygmy fruit bat.

Distribution — Philippines only; endemic to Si-
buyan Island (fmnh; Goodman & Ingle, 1993).

Habitat — Found in primary forest from near
sea level to 1325 m on Sibuyan Island, not known
from outside of primary forest (Goodman & In-
gle, 1993).

Status — Geographically restricted and subject
to pressure from deforestation (Goodman & Ingle,
1993).

Comment — Genetically distinct from H. fis-
cheri (Peterson & Heaney, 1993) and currently
being described by Goodman and Heaney.

Harpyionycteris whiteheadi Thomas,
1896. Ann. Mag. Nat. Hist., ser. 6, 18:

244.

Common Name — Harpy fruit bat.

Distribution — Philippines only, where it is
widespread, excluding the Palawan Faunal Region
and the Batanes/Babuyan group of islands. Re-
corded from Biliran (usnm), Camiguin (fmnh),
Leyte (usnm), Luzon (Camarines Sur [usnm]),
Marinduque (upd), Maripipi (ummz), Masbate
(su), Mindanao (Agusan del Norte [su], Bukidnon
[fmnh], Davao Oriental [fmnh], Lanao del Norte
[dmnh], Lanao del Sur [dmnh], Misamis Oriental
[dmnh], South Cotabato [dmnh], and Zamboanga
del Norte [dmnh] provinces), Mindoro (bmnh),
Negros (fmnh), and Samar (pnm).

Habitat — Restricted to primary or lightly dis-
turbed forest. Rare in lowland forest and moder-
ately common in montane forest to at least 1800
m. It may depend on the fruits of viney pandans
{Freycinetia spp.; Heaney, 1984; Heaney et al.,
1989; Heideman & Heaney, 1989; Rickart et al.,
1993).

Status — Populations currently stable because
of their use of montane forest, which is mostly
intact. Vulnerable to deforestation.

16

FIELDIANA: ZOOLOGY

Macroglossus minimus (E. Geoffroy,
1810). Ann. Mus. Hist. Nat. Paris, 15:97.

Common Name — Dagger-toothed flower bat,
lesser long-tongued fruit bat.

Distribution — Thailand to Australia; through-
out the Philippines. Recorded from Batu-bato
(dmnh), Biliran (usnm), Bohol (usnm), Busuanga
(usnm), Cagayan de Sulu (usnm), Calauit (ummz),
Camiguin (fmnh), Catanduanes (fmnh), Cebu
(amnh), Dinagat (usnm), Leyte (usnm), Luzon
(Albay [fmnh], Aurora [upd], Cagayan [ummz],
Camarines Norte [upd], Camarines Sur [usnm], Is-
abela [fmnh], Laguna [amnh], Quezon [ummz],
Rizal [upd], Sorsogon [fmnh], and Tarlac [usnm]
provinces), Marinduque (pnm), Maripipi (usnm),
Masbate (su), Mindanao (Bukidnon [fmnh], Da-
vao del Norte [fmnh], Davao del Sur [fmnh], Da-
vao Oriental [dmnh], Lanao del Norte [dmnh],
Lanao del Sur [dmnh], Misamis Oriental [dmnh],
South Cotabato [amnh], and Zamboanga del Sur
[dmnh] provinces), Mindoro (fmnh), Negros
(fmnh), Palawan (usnm), Panay (su), Polillo
(fmnh), Reinard (usnm), Sanga-sanga (dmnh),
Siargao (dmnh), Sibutu (dmnh), Sibuyan (fmnh,
Simunul (dmnh), and Siquijor (fmnh). Also re-
ported from Luzon (Laguna province) (Catibog-
Sinha, 1987) and from Caluya, Sibay, Semirara,
Boracay, and Carabao islands (Alcala & Alviola,
1970).

Habitat — Occurs in virtually every habitat in
the country from sea level to at least 2250 m.
Abundant in agricultural and other heavily dis-
turbed areas, common in secondary forest, and
uncommon in primary forests (Heaney et al.,
1989, in press; Heideman & Heaney, 1989; Lep-
iten, 1995; Rickart et al., 1993).

Status — Abundant and widespread, with pop-
ulations stable or increasing (Utzurrum, 1992).

Megaerops wetmorei Taylor, 1934. Mon-
ogr. Bur. Sci. Manila, 30:191.

Common Name — Mindanao fruit bat.

Distribution — Borneo to peninsular Malaysia

(Francis, 1989); in the Philippines recorded only

I on Mindanao Island (Agusan del Norte [uplb],

Bukidnon [fmnh], Davao del Sur [uplb], Davao

\ Oriental [uplb], Lanao del Norte [uplb], Lanao

I del Sur [dmnh], Misamis Oriental [uplb]. South

I Cotabato [uplb], and Zamboanga del Sur (dmnh)

provinces).

Habitat — Known in the Philippines only from

primary and lightly disturbed lowland forest from
800 m to 1200 m; probably absent from montane
and mossy forest above 1500 m (Heaney et al.,
unpubl. data). Poorly known.

Status — Within the Philippines, geographical-
ly restricted and confined to rapidly disappearing
lowland forest on Mindanao.

Nyctimene rabori Heaney and Peterson,
1984. Occas. Pap. Mus. Zool. Univ.
Michigan, 708:3.

Common Name — Philippine tube-nosed fruit
bat.

Distribution — Endemic to the Philippines; re-
corded only from Cebu (Vinciguerra & Muller,
1993), Negros (fmnh), and Sibuyan (fmnh).

Habitat — Restricted to lowland forest, usually
in primary forest but known to occur in secondary
forest on Cebu (Vinciguerra & Muller, 1993) and
Sibuyan (Goodman & Ingle, 1993). Known from
200 m to 1300 m on Negros but probably now
absent below 600 m because of the absence of
forest. Rare or uncommon at all known sites
(Heaney & Peterson, 1984; Heaney et al., 1989;
Heideman & Heaney, 1989; Mickleburgh et al.,
1992; Utzurrum, 1992). On Sibuyan, known only
from lightly disturbed mature forest near sea level
(Goodman & Ingle, 1993).

Status — Populations have declined severely
since 1950 as a result of habitat destruction, and
they face extinction on Negros Island, and per-
haps elsewhere, within 10 years if current trends
continue (Mickleburgh et al., 1992; Utzurrum.
1992). IUCN: Critically endangered.

Otopteropus cartilagonodus Kock,
Senckenberg. Biol., 50:333.

969.

Common Name — Luzon pygmy fruit bat.

Distribution — Endemic to the Philippines,
where it is widespread on Luzon Island. Recorded
from Abra (smf), Aurora (upd), Cagayan (pnm),
Camarines Sur (fmnh), Isabela (fmnh), Laguna
(pnm). Mountain (bmnh), Nueva Viscaya (pnm),
Quezon (pnm), and Zambales (pnm) provinces.

Habitat — Known only from primary and well-
developed secondary forest in lowland, montane,
and mossy forest from 200 m to 1900 m. Abun-
dance is low to moderate, usually most common
at middle elevations (Kock, 1969c, Mickleburgh

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

17

et al., 1992; Ruedas et al., 1994; Utzurrum, 1992;
Heaney et al., in press).

Status — Apparently stable because of its pri-
mary use of middle and upper elevation forest,
but poorly known. IUCN: Endangered (but we
consider the listing to be premature).

Comment — Reproductive biology documented
by Heideman et al. (1993).

Ptenochirus jagori (Peters, 1861). Mon-
atsb. K. Preuss. Akad. Wiss. Berlin, p.
707.

Common Name — Musky fruit bat.

Distribution — Endemic to the Philippines, ex-
cept the Batanes/Babuyan and Palawan faunal
regions. Specimens from Biliran (usnm), Bohol
(usnm), Bongao (dmnh), Camiguin (fmnh), Catan-
duanes (fmnh), Cebu (fmnh), Dinagat (usnm),
Leyte (usnm), Luzon (Abra [fmnh], Albay [fmnh],
Aurora [upd], Batangas [uplb], Benguet [fmnh],
Cagayan [usnm], Camarines Sur [fmnh], Isabela
[fmnh], Laguna [fmnh], Mountain Province
[fmnh], Nueva Viscaya [uplb], Pampanga [usnm],
Quezon [ummz], Rizal [uplb], Sorsogon [fmnh],
Tarlac [usnm], and Zambales [upd] provinces),
Marinduque (pnm), Maripipi (usnm), Masbate
(su), Mindanao (Agusan del Norte [dmnh], Bu-
kidnon [fmnh], Davao del Norte [fmnh], Davao
del Sur [fmnh], Davao Oriental [dmnh], Lanao del
Norte [dmnh], Lanao del Sur [dmnh], Misamis
Oriental [dmnh], Surigao del Sur [dmnh], and
Zamboanga del Sur [fmnh] provinces), Mindoro
(fmnh), Negros (fmnh), Panay (su), Polillo
(fmnh), Samar (fmnh), Sanga-sanga (dmnh), Siar-
gao (dmnh), Sibuyan (fmnh), and Siquijor (fmnh).
Also reported from Caluya, Sibay, Semirara, Bor-
acay, and Carabao islands (Alcala & Alviola,
1970).

Habitat — Abundant in primary forest, com-
mon in secondary forest, and occasionally present
in agricultural areas near forest. Ranges from sea
level to at least 1800 m, and is most common in
lowland forest, uncommon in montane forest, and
absent in mossy forest (Heaney et al., 1989, in
press; Heideman & Heaney, 1989; Ingle, 1992,
1993; Lepiten, 1995; Mudar & Allen, 1986; Rick-
art et al., 1993).

Status — Populations large and generally sta-
ble, widespread, but subject to continuing habitat
destruction (Utzurrum, 1992).

Ptenochirus minor Yoshiyuki, 1979.
Bull. Natl. Sci. Mus. Tokyo, ser. A
(Zool.), 5:75.

Common Name — Lesser musky fruit bat.

Distribution — Endemic to the Mindanao Fau-
nal Region. Specimens are from Biliran (usnm),
Dinagat (usnm), Leyte (fmnh), and Mindanao
(Agusan del Norte [dmnh], Bukidnon [fmnh],
Cotabato [amnh], and Davao del Sur [Yoshiyuki,
1979] provinces). A record from Palawan (Yoshi-
yuki, 1979) is probably erroneous.

Habitat — Recorded from sea level to 1600 m.
Common in primary lowland and montane forest
and sometimes present in mossy forest. Uncom-
mon in secondary forest (Heaney et al., 1989;
Rickart et al., 1993).

Status — Populations have declined as a result
of destruction of lowland forest habitat, but they
are still common and widespread; stable.

Pteropus dasymallus Temminck, 1825.
Monogr. Mamm. 1:180.

Common Name — Ryukyu flying fox, wooly fly-
ing fox.

Distribution — Ryukyu island group to Taiwan
and the Batanes/Babuyan region; in the Philip-
pines, recored on Batan (usnm), Dalupiri (fmnh),
and Fuga (fmnh).

Habitat — Reported to be common in forest on
the Batanes and Babuyan islands but is poorly
known (Ingle & Heaney, 1992; Utzurrum, 1992;
Ross, pers. comm.).

Status — Uncertain. Previously known popula-
tions are considered endangered to varying de-
grees (Mickleburgh et al., 1992). The Philippine
population probably is the largest for the species.
IUCN: Endangered. CITES: Appendix II.

Pteropus hypomelanus Temminck, 1853.
Esquisses Zool. sur la Cote de Guine, p.
61.

Common Name — Common island flying fox.

Distribution — Thailand to Australia. Found
throughout the Philippines except Palawan and
Batanes/Babuyan faunal regions. Records are
from Cagayan Sulu (usnm), Camiguin (dmnh),
Cebu (su), Cuyo (fmnh), Dinagat (usnm), Gui-
maras (fmnh), Leyte (ummz), Luzon (Camarines
Sur [amnh], Ilocos Norte [usnm], and Nueva Ecija

18

FIELDIANA: ZOOLOGY

[upd] provinces), Mactan (fmnh), Marinduque
(mcz), Maripipi (usnm), Negros (fmnh), Panay
(fmnh), Polillo (fmnh), Samar (usnm), Siargao
(dmnh), Sibuyan (fmnh), and Siquijor (fmnh).
Also reported from Mindanao (Gunther, 1879)
and Romblon (Timm & Birney, 1980).

Habitat — Common in agricultural areas from
sea level to ca. 900 m; absent in primary forest
(Heideman & Heaney, 1992; Rickart et al., 1993;
Utzurrum, 1992). They often roost on small is-
lands.

Status — Heavily hunted in some areas, but sta-
ble. CITES: Appendix II.

Pteropus leucopterus Temminck, 1853.
Esquisses Zool. sur la Cote de Guine, p.
60.

Common Name — Mottle-winged flying fox,
white-winged flying fox.

Distribution — Endemic to the Luzon Faunal
Region and Dinagat. Specimens have been re-
corded from Catanduanes (fmnh), Dinagat
(dmnh), and Luzon (Abra [smf], Cagayan [usnm],
Isabela [amnh], Laguna [pnm], Nueva Viscaya
[pnm], and Quezon [usnm] provinces).

Habitat — Poorly known. Moderately common
in primary montane forest on Catanduanes, pres-
ent in lowland forest (Heaney et al., 1991; Mic-
kleburgh et al., 1992; Utzurrum, 1992; Heaney et
al., unpubl. data).

Status — Poorly known but probably has de-
clined significantly as a result of habitat destruc-
tion (Utzurrum, 1992). IUCN: Endangered.
CITES: Appendix II.

Pteropus pumilus Miller,
U.S. Natl. Mus., 38:394.

1910. Proc.

Common Name — Little golden-mantled flying
fox.

Distribution — Endemic to the Philippines, ex-
cluding the Batanes/Babuyan and Palawan faunal
regions. It is also on Miangas Island, Indonesia,
adjacent to Mindanao. Records are from Balut
(usnm), Camiguin (dmnh), Leyte (dmnh), Mari-
pipi (usnm), Masbate (su), Mindanao (Zamboanga
del Sur [usnm]), Mindoro (mcz), Negros (fmnh),
Palmas (usnm), Panay (su), Sibuyan (fmnh), Si-
quijor (su), and Tablas (amnh).

Habitat — Associated with primary and well-
developed secondary lowland forest from sea lev-

el to about 1000 m. rarely to 1250 m. uncommon
outside of forest. This species is most common on
small islands and uncommon to rare on larger is-
lands (Heaney, 1984; Heaney et al., 1989; Hei-
deman & Heaney, 1989; Lepiten. 1995; Rickart et
al., 1993; Utzurrum, 1992).

Status — Declining as a result of habitat de-
struction, but still fairly widespread and stable.
IUCN: Vulnerable. CITES: Appendix II.

Comment — Includes P. balutus and P. tablasi
(Klingener & Creighton, 1984).

Pteropus speciosus K. Andersen, 1908.
Ann. Mag. Nat. Hist., ser. 8, 2:364.

Common Name — Philippine gray flying fox.

Distribution — Two islands in the Java Sea, In-
donesia, and the Mindanao and Sulu faunal
regions. Reported from Basilan (usnm), Malanipa
(usnm), Mindanao (Zamboanga del Sur Province
[fmnh]), Sanga-sanga (dmnh), Sibutu (dmnh), and
Tawi-tawi (amnh).

Habitat — Virtually unknown. Recent reports
from Tawi-tawi indicate that they are represented
by at least several large colonies.

Status — Unknown and geographically restrict-
ed. IUCN: Vulnerable. CITES: Appendix II.

Comment — Includes P. mearnsi (Heaney et al.,
1987) and may be conspecific with P. griseus
(Mickleburgh et al., 1992). Some previous reports
from Cebu, Mactan, and Negros were based on
subadult P. hypomelanus.

Pteropus vampvrus (Linnaeus, 1758).
Syst. Nat., 10th ed., 1:31.

Common Name — Large flying fox.

Distribution — Indochina to the Lesser Sundas;
throughout the Philippines except the Batanes/Ba-
buyan region. Records from Bohol (fmnh). Bon-
gao (dmnh), Cabo (usnm). Catanduanes (bmnh),
Culion (usnm), Leyte (fmnh), Guimaras (ummz),
Luzon (Abra [fmnh], Cagayan [uplb], [locos Nor-
te [usnm], Isabela [ummz]. La Union [usnm), and
Tarlac (usnm) provinces). Marinduque (pnm),
Mindanao (Agusan del Norte [UPLB], Bukidnon
[amnh], Cotabato [amnh], Davao del Norte
[fmnh], Davao del Sur [fmnh]. Davao Oriental
[uplb], Lanao del Norte [usnm], Lanao del Sur
[USNM], Maguindanao [fmnh]. Misamis Occiden-
tal [usnm], Misamis Oriental [fmnhJ, North Cot-
abato [dmnh]. South Cotabato [uplb], Zamboanga

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

19

del Notre [amnh], and Zamboanga del Sur [usnm]
provinces), Mindoro (amnh), Negros (fmnh), Pa-
lawan (fmnh), Panay (fmnh, usnm), and Tantan-
gan (usnm). Also reported from Dinagat (Gunther,
1879) and Romblon (Timm & Birney, 1980).

Habitat — Widespread and locally common in
primary lowland forest up to 1250 m; also forage
in adjacent agricultural areas (Rabor, 1955, 1986;
Rickart et al., 1993; Sanborn, 1953).

Status — Formerly occurred in many large col-
onies, but these are now greatly reduced in size
and number (Heideman & Heaney, 1989; Mickle-
burgh et al., 1992; Mudar & Allen, 1986; Rickart
et al., 1993). Heavily hunted and declining sub-
stantially (Utzurrum, 1992). CITES: Appendix II.

Pteropus sp. A.

Common Name — Mindoro pallid flying fox.

Distribution — Known only from Mindoro Is-
land (Heaney, Dans, & Crombie, unpubl. data).

Habitat — Recorded only in disturbed lowland
forest (Heaney, Dans, & Crombie, unpubl. data).

Status — Known only from patches of forest in
the lowlands of Mindoro, which have undergone
extensive deforestation (see maps in Custodio et
al., 1996; Oliver et al., 1993a). Populations cer-
tainly very small and fragmented. This species is
probably highly endangered.

Rousettus amplexicaudatus (E. Geoffroy,
1810). Ann. Mus. Hist. Nat. Paris 15:96.

Common Name — Common rousette.

Distribution — Thailand to the Solomon Is-
lands; throughout the Philippines. Records from
Balabac (usnm), Barit (fmnh), Biliran (usnm), Bo-
hol (usnm), Busuanga (usnm), Catanduanes
(fmnh), Cebu (amnh), Dalupiri (fmnh), Dinagat
(usnm), Fuga (fmnh), Jolo (amnh), Leyte (fmnh),
Lubang (mcz), Luzon (Abra [mcz], Cagayan
[ummz], Ilocos Norte [usnm], Isabela [fmnh], La-
guna [uplb], Quezon [ummz], Rizal [fmnh], Sor-
sogon [fmnh], and Zambales [usnm] provinces),
Marinduque (upd), Maripipi (usnm), Mindanao
(Bukidnon [fmnh], Davao del Norte [fmnh], Da-
vao del Sur [uplb], Lanao del Norte [upd], Lanao
del Sur [uplb], Misamis Oriental [uplb], South
Cotabato [uplb], Surigao del Sur [uplb], and
Zamboanga del Sur [mcz]), Mindoro (fmnh), Ne-
gros (fmnh), Palawan (usnm), Panay (su), Polillo
(fmnh), Samal (fmnh), Siargao (dmnh), Sibuyan

(fmnh), Siquijor (fmnh), Tablas (su), and Tincan-
san (ummz). Also reported from Caluya, Sibay,
Semirara, Boracay, and Carabao islands (Alcala
& Alviola, 1970).

Habitat — Abundant and widespread in agri-
cultural areas up to 500 m, uncommon in agri-
cultural areas from 500 m to 1100 m, and rare
elsewhere (Heaney et al., 1989, 1991, in press;
Heideman & Heaney, 1989; Lepiten, 1995; Rick-
art et al., 1993). All known roosting sites are in
caves.

Status — Locally abundant, generally stable,
but subjected to intense hunting at some cave
roosts (Utzurrum, 1992).

Emballonuridae — Sheath-tailed Bats

Three species in this family occur in the Phil-
ippines; none are endemic. A representative spe-
cies is shown in Figure 5D.

Emballonura alecto (Eydoux & Gervais,
1836). Mag. Zool. Paris, 6:7.

Common Name — Philippine sheath-tailed bat.

Distribution — Borneo, Philippines, and Sula-
wesi; probably throughout the Philippines except
the Batanes/Babuyan region. Specimens are from
Balabac (usnm), Biliran (usnm), Bohol (usnm),
Camiguin (rom), Catanduanes (usnm), Dinagat
(dmnh), Guimaras (fmnh), Leyte (usnm), Luzon
(Camarines Sur [fmnh], Laguna [cuvc], and Que-
zon [uplb] provinces), Maripipi (usnm), Minda-
nao (Davao del Norte [fmnh], Davao del Sur
[fmnh], Zamboanga del Norte [fmnh], and Zam-
boanga del Sur [amnh] provinces), Negros
(fmnh), and Panay (usnm).

Habitat — Recorded only in lowland areas (450
m and below) in disturbed forest and agricultural
areas with scattered remnant forest. Most records
are from individuals captured in caves, under
large boulders, or in man-made tunnels (Heaney
et al., 1991, in press; Ingle, 1992; Rabor, 1986;
Rickart et al., 1993; Taylor, 1934).

Status — Common in areas with caves in or
near forest; apparently rare elsewhere.

Saccolaimus saccolaimus (Temminck,
1838). Tijdschr. Nat. Gesch. Physiol., 5:
14.

Common Name — Pouched bat.

Distribution — Widespread, from India to Ti-

20

FIELDIANA: ZOOLOGY

mor, New Guinea, and northern Australia. In the
Philippines, recorded from Catanduanes (usnm),
Mindanao (Misamis Oriental [dmnh] and Zam-
boanga del Sur [usnm] provinces), and Negros
(usnm).

Habitat — Recorded from sea level to 800 m.
Poorly known; may be moderately common in ag-
ricultural areas.

Status — Unknown.

Comment — Philippine population formerly
considered to be a distinct species, S. pluto (Cor-
bet & Hill, 1992; Heaney et al., 1991; Koopman,
1993).

Taphozous melanopogon Temminck,
1841. Monogr. Mamm., 2:287.

Common Name — Black-bearded tomb bat.

Distribution — Widespread from Sri Lanka to
Southeast Asia and the Lesser Sunda Islands. Re-
corded from Biliran (ummz), Cebu (amnh), Gi-
gante (ummz), Leyte (ummz), Luzon (Ilocos Norte
[usnm], Nueva Viscaya [uplb], Pangasinan
[fmnh], and Rizal [usnm] provinces), Maripipi
(usnm), Mindanao (Davao del Sur [fmnh]), Ne-
gros (fmnh), Palawan (ummz), Sibuyan (fmnh),
and Tincasan (ummz). Also reported from Lubang,
Luzon (Abra Province) and Mindoro (Lawrence,
1939).

Habitat — Common in urban areas, in areas
with limestone caves, and in sea caves, at eleva-
tions at or below 150 m (Lawrence, 1939; Rickart
et al., 1993; Sanborn, 1952; Taylor, 1934). Often
roost in shallow portions of caves; and in church-
es and other buildings with large attics.

Status — Widespread in Asia; abundant in the
Philippines.

Comment — Philippine populations were for-
merly separated as T. philippinensis. We consider
the latter to be a subspecies of this widespread
species (Corbet & Hill, 1992; Koopman, 1993;
Rickart et al., 1993).

Megadermatidat
Bats

-False Vampire and Ghost

Megaderma spasma (Linnaeus, 1758).
Syst. Nat., 10th ed., 1:32.

Common Name — Common Asian ghost bat.
Lesser false vampire.

Distribution — India to the Molucca Islands;
throughout the Philippines except the Batanes/Ba-
buyan region. Recorded on Biliran (usnm), Bohol
(usnm), Busuanga (usnm), Catanduanes (usnm),
Cebu (amnh), Dinagat (dmnh), Leyte (usnm), Lu-
zon (Aurora [upd], Camarines Sur [usnm], Isabela
[fmnh], and Rizal [usnm] provinces), Mindanao
(Davao del Norte [fmnh], Lanao del Norte
[usnm], Misamis Occidental Iusnm], South Cota-
bato [fmnh], and Zamboanga del Sur [uplb] prov-
inces), Mindoro (fmnh), Negros (fmnh), Palawan
(ummz), Panay (su), Polillo (fmnh), and Siquijor
(su). Also reported from Davao Oriental Province
(Mindanao) and Negros by Sanborn (1952), from
Abra Province (Luzon) and Cebu by Lawrence
(1939), and from Laguna Province (Luzon) by
Taylor (1934) and Ingel (1992).

Habitat — Lowland primary and disturbed for-
est from sea level to 950 m (Heaney et al., 1991).
Known to roost in caves, tree-hollows, and hollow
logs (Heaney et al., in press; Ingle, 1992; Law-
rence, 1939; Lepiten, 1995; Rabor, 1986; Rickart
et al., 1993; Taylor, 1934).

Status — Widepread, locally common to un-
common in primary forest and secondary forest.

Rhinolophidac
Bats

-Horseshoe and Roundleaf

A single species from this family occurs in the
Philippines; its appearance is shown in Figure 5B.

With 18 species currently recognized in the
Philippines, this is one of the most diverse fami-
lies of mammals in the Philippines; the appear-
ance of a representative species is shown in Fig-
ure 5C. However, the family is poorly known,
principally because rhinolophids are difficult to
capture with mist nets. The dearth of museum
specimens (both from the Philippines and else-
where) is a major reason for continuing uncer-
tainty regarding the taxonomy of several members
of this family (Hill, 1983). We expect future sur-
veys, especially those employing harp traps, to
add to both the number and known distribution of
the rhinolophids from the Philippines. Many spe-
cies require caves as roosting sites and have been
impacted negatively by the widespread distur-
bance of caves. Other species roost in large hol-
low trees, especially in lowland dipterocarp forest,
and have been severely affected by logging that

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

21

destroys both the roosting trees and the foraging
habitat.

Coelops hirsutus (Miller, 1911). Proc.
U.S. Natl. Mus., 38:395.

Common Name — Philippine tailless roundleaf
bat.

Distribution — Specimens only from Mindanao
(Bukidnon [fmnh]) and Mindoro (usnm).

Habitat — May be dependent on caves.

Status — Unknown.

Comment — May be a subspecies of C. robin-
soni, known from the Malay Peninsula and Bor-
neo (Corbet & Hill, 1992).

Hipposideros ater Templeton, 1848. J.
Asiat. Soc. Bengal, 17:252.

Common Name — Dusky roundleaf bat.

Distribution — India to Australia; throughout
the Philippines. Recorded from Bohol (usnm), Ca-
tanduanes (usnm), Cebu (amnh), Leyte (usnm),
Luzon (Abra [smf], Cagayan [fmnh], Camarines
Sur [usnm], Laguna [fmnh], Pampanga [usnm],
Rizal [usnm], and Tarlac [usnm] provinces), Mar-
ipipi (usnm), Mindanao (Davao del Sur [fmnh]),
Mindoro (fmnh), Negros (fmnh), and Palawan
(fmnh). Also reproted from Balabac by Hill
(1963), Benguet Province (Luzon) by Taylor
(1934), and Marinduque and Mindoro by Law-
rence (1939).

Habitat — Poorly known; recorded from sea
level to 1200 m in lowland and montane forest.
Known to roost in caves in forest and in agricul-
tural areas (Heaney et al., 1991; Rickart et al.,
1993).

Status — Geographically widespread. In the
Philippines, probably has declined as a result of
destruction of lowland forest and distrubance of
caves.

Comment — Hill (1963) included H. wrighti
Taylor 1934 as a synonym of this species.

Hipposideros bicolor (Temminck, 1834).
Tijdschr. Nat. Gesch. Physiol., 1:19.

Common Name — Bicolored roundleaf bat.

Distribution — India to Timor; Philippine spec-
imens from Luzon (Camarines Sur Province
[usnm]), Mindoro (mcz), and Palawan (ummz).

Habitat — Unknown in the Philippines. Roosts
in caves in peninsular Malaysia (Payne et al.,
1985).

Status — Unknown.

Hipposideros cervinus (Gould, 1863).
Mamm. Austr., 3: pi. 34.

Common Name — Fawn-colored roundleaf bat.

Distribution — Peninsular Malaysia to Austra-
lia and Vanuatu; Philippine specimens from Min-
danao (Davao del Sur [fmnh] and Maguindanao
[fmnh]) only.

Habitat — Virtually unknown in the Philip-
pines; recorded in caves on Mindanao (Sanborn,
1952). On Borneo, usually roosts in caves and for-
ages in forest (Payne et al., 1985).

Status — Unknown.

Comment — The taxonomic status and distribu-
tion of this species were discussed by Jenkins and
Hill (1981).

Hipposideros coronatus (Peters, 1871).
Monatsb. K. Preuss. Akad. Wiss. Berlin,
p. 327.

Common Name — Large Mindanao roundleaf
bat.

Distribution — Known only from Mainit, Su-
rigao del Norte Province, Mindanao (Peters,
1871).

Habitat — Unknown.

Status — Unknown; geographically restricted.

Hipposideros diadema (E. Geoffroy,
1813). Ann. Mus. Hist. Nat. Paris, 20:
263.

Common Name — Diadem roundleaf bat.

Distribution — Burma to the Solomon Islands;
throughout the Philippines except the Batanes/Ba-
buyan region. Specimens from Bohol (usnm),
Busuanga (usnm), Calauit (ummz), Catanduanes
(usnm), Dinagat (dmnh), Guimaras (fmnh), Leyte
(usnm), Luzon (Bulacan [usnm], Cagayan [pnm],
Camarines Sur [usnm], Ilocos Norte [usnm], Isa-
bela [fmnh, ummz], Laguna [amnh], Nueva Viz-
caya [ummz], Pampanga [usnm], and Quezon
[ummz] provinces), Mindanao (Agusan del Norte
[su], Bukidnon [fmnh], Davao Oriental [uplb],
Lanao del Norte [uplb], Maguindanao [fmnh],

22

FIELDIANA: ZOOLOGY

and Zamboanga del Norte [su]), Negros (fmnh),
Palawan (usnm), Panay (su), Polillo (fmnh), Sa-
mar (usnm), and Siquijor (fmnh). Also reported
from South Cotabato Province (Mindanao) by
Sanborn (1952), from Cebu and Mindoro by Law-
rence (1939), from Benguet and Rizal provinces
(Luzon), and Zamboanga del Sur Province (Min-
danao) by Taylor (1934).

Habitat — In primary forest and disturbed low-
land areas from sea level to 900 m. Roost in hol-
low trees and in caves and human-made tunnels
(Heaney et al., in press; Lepiten, 1995; Rickart et
al., 1993; Sanborn, 1952).

Status — Widespread and common.

Hipposideros lekaguli Thonglongya and
Hill, 1974. Mammalia, 38:286.

Common Name — Large Asian roundleaf bat.

Distribution — Thailand, peninsular Malaysia,
and the Philippines. In the Philippines, recorded
only on Luzon (Isabela [fmnh] Province) and
Mindoro (fmnh) (Balete et al., 1995).

Habitat — The two known specimens from the
Philippines were collected close to sea level in
ultrabasic forest and in an agricultural area near a
river (Balete et al., 1995).

Status — Unknown.

Comment — As currently defined, this species is
variable geographically. A comprehensive revi-
sion is needed.

Hipposideros obscurus (Peters, 1861).
Monatsb. K. Preuss. Akad. Wiss. Berlin,
p. 707.

Common Name — Philippine forest roundleaf
bat.

Distribution — Philippines only. Specimens are
from Bohol (usnm), Catanduanes (fmnh), Dinagat
(dmnh), Luzon (Benguet [amnh], Camarines Sur
[usnm], Laguna [amnh], Pampanga [usnm], and
Tarlac [usnm] provinces), Maripipi (usnm), Min-
danao (Bukidnon [fmnh] and South Cotabato
[amnh] provinces), Negros (usnm), and Siquijor
(su).

Habitat — Locally common to uncommon in
primary and disturbed forest up to 850 m (Heaney
et al., in press; Lepiten, 1995; Rickart et al.,
1993). Several records of specimens taken in
caves, one in a mine shaft, one in a dark cavity

in a tree buttress (Heaney et al.. 1991; Taylor,
1934), and one in a hollow tree (Sanborn. 1952).
Status — Widespread, but dependent on forest
and perhaps on caves.

Hipposideros pygmaeus (Waterhouse,
1843). Proc. Zool. Soc. Lond. p. 67.

Common Name — Philippine pygmy roundleaf
bat.

Distribution — Philippines only. Specimens
taken from Bohol (usnm), Luzon (Camarines Sur
[fmnh], and Rizal [usnm] provinces), Marinduque
(pnm), Negros (fmnh), and Panay (su).

Habitat — The few specimens were taken in
caves from sea level to 200 m, in or near forest
(Sanborn, 1952; Taylor, 1934).

Status — Widespread but seemingly rare. This
species is probably strongly impacted by the de-
struction of cave habitats.

Rhinolophus acuminatus Peters, 1871.
Monatsb. K. Preuss. Akad. Wiss. Berlin,
p. 308.

Common Name — Acuminate horseshoe bat.

Distribution — Thailand to Lombok; in the
Philippines, known only from the Palawan Faunal
Region. Recorded from Balabac (usnm), Busuan-
ga (usnm), and Palawan (fmnh).

Habitat — Unknown in the Philippines. Occurs
in lowland dipterocarp forest on Borneo (Payne
et al., 1985).

Status — Uncertain, apparently locally com-
mon.

Rhinolophus anderseni Cabrera, 1909.
Bol. Real. Soc. Esp. Hist. Nat., p. 305.

Common Name — Andersen's horseshoe bat.

Distribution — Philippines only, where it is re-
corded from Luzon (Cabrera, 1909) and Palawan
(Allen, 1922; ummz).

Habitat — Unknown.

Status — Unknown, rare in collections.

Comment — Corbet and Hill (1992) consider
this species close to R. nereis and R. borneensis.
The form R. anderseni aequalis Allen 1922.
which is the basis for the Palawan record, is of
uncertain relationship. We consider this group to
be badly in need of review.

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

23

Rhinolophus arcuatus Peters, 1871.
Monatsb. K. Preuss. Akad. Wiss. Berlin,
p. 305.

Common Name — Arcuate horseshoe bat.

Distribution — Sumatra to New Guinea;
throughout the Philippines, possibly excluding the
Palawan Faunal Region. Specimens from Biliran
(usnm), Camiguin (dmnh), Catanduanes (usnm),
Dalupiri (fmnh), Fuga (fmnh), Guimaras (usnm),
Leyte (usnm), Luzon (Abra [smf], Cagayan [pnm],
Camarines Sur [fmnh], Isabela [fmnh], Laguna
[fmnh], Pampanga [usnm], and Rizal [usnm] prov-
inces), Maripipi (usnm), Masbate (su), Mindanao
(Bukidnon [fmnh], Davao del Sur [fmnh], Ma-
guindanao [fmnh], Zamboanga del Norte [fmnh],
and Zamboanga del Sur [bmnh] provinces), Min-
doro (fmnh), Negros (fmnh), Panay (su), Polillo
(fmnh), Sibutu (dmnh), Sibuyan (fmnh), Siquijor
(su), and Tawi-tawi (dmnh).

Habitat — From lowlands to at least 1050 m, in
agricultural lands to primary forest (Heaney et al.,
1991, in press; Lepiten, 1995; Rickart et al.,
1993). Roosting sites sometimes in caves (San-
born, 1952.)

Status — Widespread, locally common.

Comment — Heaney et al. (1991), Ingle and
Heaney (1992), and Rickart et al. (1993) have
commented that, on most islands, there appears to
be a smaller lowland morph associated with caves
in agricultural areas and a larger highland morph
associated with primary forest. We have also not-
ed subtle but consistent differences between pop-
ulations on each Pleistocene island. Further sys-
tematic studies are needed.

Status — Locally abundant, but dependent on
primary forest.

Comment — As noted by Ingle and Heaney
(1992), there are several problems with the tax-
onomy of R. inops. First, the holotype of R. inops
has a distinctively shaped nose-leaf not found in
any of the specimens referred to this species. Sec-
ond, there is considerable geographic variation,
and species as currently defined may represent a
species group. Careful study is needed.

Rhinolophus macrotis Blyth, 1844. J.
Asiat. Soc. Bengal, 13:485.

Common Name — Big-eared horseshoe bat.

Distribution — India to Sumatra and the Phil-
ippines. Specimens from Guimaras (usnm), Luzon
(Abra [smf] and Pampanga [usnm]), Mindanao
(Bukidnon [smf]), and Negros (fmnh).

Habitat — Recorded in lowland forest from 200
m to 1050 m. Poorly known in the Philippines.
There are some records from caves in forest.

Status — Widespread but seemingly uncom-
mon.

Comment — The Philippine form of R. macrotis
was initially described as a separate species, R.
hirsutus (Andersen, 1905), but was later sub-
sumed under R. macrotis by Tate (1943). Philip-
pine populations are morphologically distinct
from all others and may deserve recognition as a
distinct species (Ingle & Heaney, 1992). More
specimens are needed to assess this issue.

Rhinolophus inops K. Andersen, 1905.
Ann. Mag. Nat. Hist., ser. 7, 16:284, 651.

Common Name — Philippine forest horseshoe
bat.

Distribution — Philippines only, where it is re-
corded from Biliran (usnm), Camiguin (msu-iit),
Catanduanes (usnm), Leyte (usnm), Luzon (Ca-
marines Sur [usnm], and Pampanga [usnm] prov-
inces), Mindanao (Bukidnon [fmnh] and Davao
del Sur [usnm] provinces), Negros (fmnh), and
Polillo (fmnh).

Habitat — Common to abundant in primary
lowland and montane forest from sea level to
2250 m, rarely in secondary forest and mossy for-
est (Heaney et al., 1991, in press; Rickart et al.,
1993).

Rhinolophus philippinensis Waterhouse,
1843. Proc. Zool. Soc. Lond., p. 68.

Common Name — Enormous-eared horsehoe
bat.

Distribution — Borneo and the Philippines to
Australia (Tate, 1943); Philippine records from
Luzon (Abra Province [smf]), Mindanao (Zam-
boanga del Norte [fmnh], and Zamboanga del Sur
[uplb] provinces), Mindoro (mcz), Negros (fmnh),
and Siquijor (fmnh).

Habitat — Recorded in primary and secondary
forest from 200 m to 1500 m (Lepiten, 1995; Rue-
das et al., 1994; Heaney, unpubl. data).

Status — Uncertain, but uncommon. Captured
only in forest.

24

FIELDIANA: ZOOLOGY

Rhinolophus rufus Eydoux and Gervais,
1836. In Laplace, Voy. autour du monde
par les mers de l'lnde .... la Favorite,
5(Zoologie), part 2:9.

Common Name — Large rufous horseshoe bat.

Distribution — Philippines only, where it is re-
corded from Bohol (usnm), Catanduanes (usnm),
Leyte (dmnh), Luzon (Laguna [amnh], Pampanga
[usnm], and Rizal [usnm] provinces), Marinduque
(fmnh), Mindanao (Davao del Sur [fmnh] and
Maguindanao [fmnh] provinces), Mindoro (mcz),
Polillo (fmnh).

Habitat — Recorded in primary and good sec-
ondary forest, either in or near caves (Heaney et
al., 1991; Lawrence, 1939; Sanborn, 1952).

Status — Uncertain; generally uncommon and
probably dependent on lowland caves, most of
which have been heavily disturbed.

from Mindanao (South Cotabato Province [Ta\ lor
1934]) and Lubang (Lawrence, 1939).

Habitat — Primary lowland forest from 250 m
to 1 100 m, with several records from caves (Hea-
ney et al.. 1991; Rickart et al., 1993) and one from
a darkened cavity in a tree buttress (Taylor, 1934).

Status — Widespread and moderately common.

Comment — Closely related to R. celebensis
(Corbet & Hill, 1992).

Vespertilionidae — Vesper and Evening Bats

Twenty-two species of vespertilionids, includ-
ing the recently recorded Harpiocephalus harpia
(Rickart et al., 1993), are now known from the
Philippines. The appearance of a representative
species is shown in Figure 5F. Only one species,
Myotis rufopictus, is considered to be endemic.

Rhinolophus subrufus K. Andersen,
1905. Ann. Mag. Nat. Hist., ser. 7, 16:
283.

Common Name — Small rufous horseshoe bat.

Distribution — Philippines only; specimens
from Camiguin (dmnh), Catanduanes (usnm), Lu-
zon (Abra [fmnh], Camarines Sur [usnm], Isabela
[fmnh], Laguna [fmnh], Pampanga [usnm], Rizal
[bmnh], and Sorsogon [fmnh] provinces); Min-
danao (Davao del Sur [usnm] and South Cotabato
[amnh] provinces), and Mindoro (fmnh).

Habitat — Poorly known. Recorded from near
sea level to over 1000 m, with some records from
caves.

Status — Unknown. IUCN: Vulnerable. We be-
lieve this listing to be premature, given the current
lack of knowledge.

Rhinolophus virgo K. Andersen, 1905.
Proc. Zool. Soc. Lond. p. 88.

Common Name — Yellow-faced horseshoe bat.

Distribution — Philippines only, found through-
out the Philippines. Records from Batan (usnm),
Busuanga (usnm), Catanduanes (usnm), Cebu
(amnh), Jolo (amnh), Leyte (usnm), Luzon (Abra
[smf], Camarines Sur [usnm], Isabela [fmnh], La-
guna [amnh], Pampanga [usnm], and Sorsogon
[fmnh] provinces), Maripipi (usnm), Mindanao
(Bukidnon Province [smf]), Negros (fmnh), Pa-
lawan (fmnh), and Sibuyan (fmnh). Also reported

Glischropus tylopus (Dobson, 1875).
Proc. Zool. Soc. Lond., p. 473.

Common Name — Thick-thumbed pipistrelle.

Distribution — Burma to Molucca Islands;
Philippine records from Palawan only (usnm).

Habitat — Unknown in the Philippines. Roosts
in rock crevices, in hollow bamboo, and in new
banana leaves in peninsular Malaysia (Payne et
al., 1985).

Status — Unknown.

Harpiocephalus harpia (Temminck,
1840). Monogr. Mamm., 2:219.

Common Name — Hairy-winged bat.

Distribution — India to Indochina and Taiwan.
Java, Molucca, and the Sunda Islands. In the Phil-
ippines recorded only from Luzon (Camarines Sur
[usnm] Province), Leyte (usnm), Negros (fmnh),
and Panay (pawb and Japan Wildlife Research
Center).

Habitat — In the Philippines, known from pri-
mary and disturbed lowland forest from 475 m to
750 m (Heaney et al., in press; Ingle & Heaney.
1992; Rickart et al., 1993; Utzurrum. unpubl.
data).

Status — Widespread in southern Asia; proba-
bly dependent on lowland forest.

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

25

Kerivoula hardwickii (Horsfield, 1824).
Zool. Res. Java, part 8:28.

Common Name — Common woolly bat.

Distribution — India and southern China to
Lesser Sunda Islands. Philippine records are from
Biliran (usnm), Leyte (usnm), Luzon (Camarines
Sur [usnm] Province), Mindanao (Bukidnon
[fmnh] Province), and Palawan (ummz). Also re-
ported from Samar (Taylor, 1934).

Habitat — In the Philippines, recorded from
500 m to 1600 m in lowland, montane, and
ridgetop mossy forest (Heaney et al., in press;
Rickart et al., 1993).

Status — Widespread in southern Asia. Proba-
bly moderately common in primary forest.

Kerivoula pellucida (Waterhouse, 1845).
Proc. Zool. Soc. Lond., p. 6.

Common Name — Clear-winged woolly bat.

Distribution — Borneo, Java, Malay Peninsula,
Sumatra, and the Philippines; records are from
Mindanao (Davao del Norte [fmnh]) and Palawan
(ummz). Also reported from Jolo (Taylor, 1934).

Habitat — Taylor (1934) reported two groups
of this species that were roosting in dead leaves
on small shrubs in lowland forest.

Status — Unknown.

Kerivoula whiteheadi Thomas, 1894.
Ann. Mag. Nat. Hist., ser. 6, 14:460.

Common Name — Whitehead's woolly bat.

Distribution — Southern Thailand to Borneo
and the Philippines; records are from Luzon (Is-
abela Province [bmnh]), Mindanao (Davao del
Norte [fmnh], Davao del Sur [fmnh], and Zam-
boanga del Norte [fmnh] provinces), and Palawan
(ummz). Also reported from Mindanao (Davao del
Sur) by Sanborn (1952) and Panay by Taylor
(1934).

Habitat — Known only from near sea level, in
disturbed forest and agricultural areas (Sanborn,
1952).

Status — Unknown.

Miniopterus australis Tomes, 1858. Proc.
Zool. Soc. Lond. p. 125.

Common Name — Little bent-winged bat.
Distribution — India to Australia and through-

out the Philippines except the Babuyan/Batanes
group. Specimens from Bongao (uplb), Bohol
(usnm), Capiz (usnm), Catanduanes (usnm), Gui-
maras (fmnh), Leyte (usnm), Luzon (Bulacan
[usnm] and Rizal [usnm] provinces), Mindanao
(Davao del Norte [fmnh], Davao del Sur [fmnh],
Maguindanao [fmnh], and Zamboanga del Norte
[fmnh]), Negros (fmnh), Panay (fmnh), Polillo
(fmnh), and Siquijor (su).

Habitat — Roosts in caves in lowland areas
from sea level to about 200 m in agricultural areas
or second growth (Heaney et al., 1991; Lepiten,
1995; Rickart et al., 1993; Sanborn, 1952; Taylor,
1934).

Status — Geographically widespread and com-
mon but dependent on caves.

Comment — Of the three size classes of bats of
this genus that we have examined from the Phil-
ippines, all of the smallest class can be referred
to a single species; all appear to be this species,
rather than M. pusillus (Koopman, 1993).

Miniopterus schreibersi (Kuhl, 1817).
Die Deutschen Fledermause, Hanau, p.
14.

Common Name — Common bent-winged bat.

Distribution — Europe to the Solomon Islands;
throughout the Philippines. Specimens from Bo-
hol (usnm), Catanduanes (usnm), Dalupiri (fmnh),
Guimaras (fmnh), Leyte (usnm), Luzon (Cagayan
[ummz], Camarines Sur [fmnh], Isabela [ummz],
Pampanga [usnm], Rizal [usnm], and Tarlac
[usnm] provinces), Mindanao (Bukidnon [fmnh],
Davao del Sur [fmnh], and Zamboanga del Norte
[fmnh] provinces), Negros (fmnh), Panay (fmnh),
and Polillo (fmnh). Also reported from Cebu, Lu-
zon (Benguet), Marinduque, Mindoro, and Tablas
(Hollister, 1913; Lawrence, 1939; Sanborn, 1952;
Taylor, 1934).

Habitat — Common in caves throughout the
lowlands in agricultural areas and in forest. Netted
in primary forest from near sea level to 1450 m
in lowland and montane forest; occasionally also
use man-made tunnels (Heaney et al., 1991, in
press; Lawrence, 1939; Rickart et al., 1993; San-
born, 1952; Taylor 1934).

Status — Common and widespread, dependent
on caves.

26

FIELDIANA: ZOOLOGY

Miniopterus tristis (Waterhouse, 1845).
Proc. Zool. Soc. Lond., p. 3.

Common Name — Greater bent-winged bat.

Distribution — Philippines to Solomon Islands;
records throughout the Philippines, possibly ex-
cluding the Palawan Faunal Region. Recorded
from Bohol (rom), Guimaras (rom), Leyte
(usnm), Lubang (usnm), Mindanao (Maguindanao
[fmnh] and Zamboanga del Norte [rom] prov-
inces), Negros (fmnh), and Tablas (rom). Also re-
ported from Cebu, Lubang, Luzon, Mindoro, and
Samar (Hollister, 1912, 1913; Lawrence, 1939;
Peterson, 1981; Taylor, 1934).

Habitat — Apparently roost only in caves and
forage in agricultural areas and disturbed lowland
forest near sea level (Rickart et al., 1993; San-
born, 1952).

Status — Moderately common and widespread,
but dependent on caves.

Murina cyclotis Dobson, 1872. Proc.
Asiat. Soc. Bengal, p. 210.

Common Name — Round-eared tube-nosed bat.

Distribution — Sri Lanka to Hainan and Bor-
neo. In the Philippines, recorded from Biliran
(ummz), Camiguin (msu-iit), Catanduanes (usnm),
Luzon (Camarines Sur [usnm] Province), Minda-
nao (Bukidnon [fmnh] Province), Sibuyan (fmnh),
and Siquijor (su).

Habitat — Primary lowland, lightly disturbed
lowland, and lower montane forest from 250 m to
1500 m (Heaney et al., 1991, in press; Lepiten,
1995; Rickart et al., 1993; Ruedas et al., 1994).

Status — Widespread and moderately common
in the Philippines.

Myotis horsefieldii (Temminck,
Monogr. Mamm., 2:226.

1840).

Common Name — Common Asiatic myotis.

Distribution — Southeastern China to the Ma-
lay Peninsula, Bali, and Sulawesi. Philippine re-
cords are from Bohol (usnm), Catanduanes
(fmnh), Luzon (Cagayan [fmnh], Laguna [cuvc],
Pampanga Iusnm], Quezon [ummz], and Rizal
[usnm] provinces), Mindanao (Lanao del Norte
Province [dmnh]), Negros (ummz), and Palawan
(ummz).

Habitat — In the Philippines, recorded in low-
land forest and in agricultural areas from sea level

to 800 m; sometimes roosts in caves ami n tun-
nels and reported roosting beneath a large i\>ck
over a stream (Taylor, 1934).

Status — Widespread and moderately common;
may depend on caves.

Comment — Includes M. jeannei (Hill, 1983).

Myotis macrotarsus (Waterhouse, 1845).
Proc. Zool. Soc. Lond. p. 5.

Common Name — Philippine large-footed my-
otis.

Distribution — Borneo and the Philippines;
probably throughout the Philppines. Specimens
are from Guimaras (fmnh), Marinduque (fmnh),
Mindanao (Lanao del Norte [usnm] Province),
Negros (fmnh), Palawan (anm), and Polillo
(fmnh). Also reported from Luzon (Rizal Prov-
ince), Marinduque, Mindanao (Zamboanga), and
Tawi-tawi (Hollister, 1913; Lawrence, 1939; Tay-
lor, 1934).

Habitat — Roosts only in caves near sea level,
forages in agricultural areas (Heaney & Utzurrum,
unpubl. data).

Status — Uncommon, dependent on caves.

Myotis muricola (Gray, 1846). Cat.
Hodgson Coll. Brit. Mus., p. 4.

Common Name — Whiskered myotis.

Distribution — Afghanistan to New Guinea;
throughout the Philippines. Recorded from Biliran
(ummz), Busuanga (fmnh), Culion (fmnh), Leyte
(usnm), Luzon (Camarines Sur [usnm], Laguna
[amnh], and Rizal [amnh] provinces), Mindanao
(Bukidnon [fmnh] and South Cotabato [amnh]),
Maripipi (usnm), and Negros (fmnh).

Habitat — In primary and secondary lowland
and montane forest from near sea level to 1 125 m
(Rickart et al., 1993; Heaney et al., in press, un-
publ. data).

Status — Common and widespread in Asia.

Comment — Includes M. browni, M. herrei, and
M. patriciae (Koopman. 1993).

Myotis rufopictus (Waterhouse, 1845).
Proc. Zool. Soc. Lond., p. 3, 8.

Common Name — Orange -fingered myotis.
Distribution — Philippines only; recorded on
Luzon (Bulacan [usnm], Camarines Sur [usnm],

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

27

Isabela [ummz], Nueva Viscaya [uplb], and Rizal
[fmnh] provinces), Negros (umnz), Palawan
(ummz), and Sibuyan (fmnh).

Habitat — Recorded in primary lowland and
montane forest from 50 m to 1125 m and in an
agricultural area near sea level (Mudar & Allen,
1986; Heaney et al., in press, unpubl. data).

Status — Unknown, probably uncommon.

Comment — Considered to be a subspecies of
M. formosus by Corbet and Hill (1992) and Koop-
man (1993), but we disagree on the basis of cra-
nial and external morphology.

Philetor brachypterus (Temminck,
1840). Monogr. Mamm., 2:215.

Common Name — Short-winged pipistrelle.

Distribution — Nepal to New Guinea; through-
out most of the Philippines. Recorded from Ca-
tanduanes (fmnh), Leyte (usnm), Luzon (Camari-
nes Sur [usnm] and Laguna [cuvc]), Mindanao
(Bukidnon [fmnh], Misamis Oriental [fmnh], and
Zamboanga del Sur [dmnh]) provinces), and Ne-
gros (ummz) (Kock, 1981).

Habitat — In primary and disturbed lowland
forest from 475 m to 900 m (Rickart et al., 1993;
Heaney et al., in press, unpubl. data).

Status — Uncertain; probably moderately com-
mon in primary forest.

[fmnh] provinces), Mindanao (Bukidnon [fmnh]
and Davao del Sur [fmnh] provinces), Mindoro
(fmnh), Negros (fmnh), Palawan (fmnh), Panay
(usnm), Reinhard (ummz), and Sibuyan (fmnh).

Habitat — Common in primary montane forest,
uncommon in primary lowland and mossy forest,
from sea level to 1750 m (Sanborn, 1952; Heaney
et al., in press, unpubl. data).

Status — Moderately common and widespread
in eastern Asia.

Comment — Previous reports of P. imbricatus
from the Philippines all appear to be based on
specimens of this species.

Pipistrellus petersi (Meyer, 1899). Abh.
Zool. Anthrop. -Ethnology. Mus. Dres-
den, 7(7): 13.

Common Name — North Wallacean pipistrelle.

Distribution — Sulawesi, Molucca Islands, and
the Philippines; records from Luzon (Benguet
Province [usnm]) and Mindanao (Davao del Sur
[fmnh] and Surigao del Norte [amnh] provinces).

Habitat — Unknown.

Status — Unknown.

Pipistrellus stenopterus (Dobson, 1875).
Proc. Zool. Soc. Lond. p. 470.

Phoniscus jagori (Peters, 1866). Mon-
atsb. K. Preuss. Akad. Wiss. Berlin, p.
399.

Common Name — Common trumpet-eared bat.

Distribution — Bali, Borneo, Java, Sulawesi,
and Samar.

Habitat — Unknown in Philippines.

Status — Unknown.

Comment — Koopman (1993) considered Phon-
iscus to be a subgenus of Kerivoula.

Pipstrellus javanicus (Gray, 1838). Mag.
Zool. Bot., 2:498.

Common Name — Javan pipistrelle.

Distribution — Korea to Java and the Philip-
pines; throughout the Philippines. Specimens
from Camiguin (fmnh), Luzon (Benguet [fmnh],
Cagayan [fmnh], Camarines Sur [fmnh], Isabela
[fmnh], Laguna [fmnh], Quezon [umz], and Rizal

Common Name — Narrow-winged pipistrelle.

Distribution — Sumatra to Mindanao; the sin-
gle Philippine specimen is from Mindanao (Zam-
boanga del Sur Province [usnm]).

Habitat — Unknown in the Philippines; report-
ed to roost in houses and feed over open fields on
Borneo (Payne et al., 1985).

Status — Unknown.

Pipistrellus tenuis (Temminck, 1840).
Monogr. Mamm., 2:229.

Common Name — Least pipistrelle.

Distribution — Thailand to Australia; Philip-
pine records from Luzon (Rizal [usnm]), Negros
(fmnh), and Sibuyan (fmnh). Also reported from
Mindanao by Taylor (1934).

Habitat — Primary lowland and montane forest
from 800 m to 1700 m (Heaney et al., unpubl.
data).

Status — Widespread and moderately common.

28

FIELDIANA: ZOOLOGY

Scotophilus kuhlii Leach, 1821. Trans.
Linn. Soc. Lond. 13:71.

Common Name — Lesser Asian house bat.

Distribution — Pakistan to Taiwan and Bali.
This species is found throughout the Philippines.
Recorded from Biliran (usnm), Catanduanes
(usnm), Cuyo (fmnh), Guimaras (usnm), Leyte
(fmnh), Luzon (Bulacan [fmnh], Cagayan [ummz],
Cavite [usnm], Isabela [fmnh], Laguna [uplb]. La
Union [usnm], Pampanga [fmnh], Rizal [fmnh],
and Zambales [usnm] provinces), Maripipi
(usnm), Mindanao (Davao del Sur [fmnh], Lanao
del Norte [usnm], Maguindanao [fmnh], Misamis
Oriental [uplb], and South Cotabato [fmnh] prov-
inces), Negros (fmnh), Palawan (fmnh), Panay
(fmnh), Sibuyan (fmnh), and Ticao (usnm). Also
reported from Cebu and Luzon (Abra) (Lawrence,
1939; Taylor, 1934). Also reported from Carabao
Island (Alcala & Al viola, 1970).

Habitat — Commonly roost in buildings and in
"tents" formed from modified palm fronds. They
forage in urban and agricultural areas and second-
ary forest from sea level to about 600 m (Rickart
et al., 1989b, 1993; Heaney et al., unpubl. data).

Status — Abundant in urban and agricultural
areas.

Tvlonycteris pachypus (Temminck,
1840)." Monogr. Mamm., 2:217.

Common Name — Lesser flat-headed bat, Lesser
bamboo bat.

Distribution — India to the Philippines and
Lesser Sunda Islands; probably throughout the
Philippines. Recorded from Calauit (ummz), Cu-
lion (fmnh), Luzon (Rizal Province [usnm]), and
Palawan (usnm).

Habitat — Bamboo stands in lowland agricul-
tural areas.

Status — Widespread, probably moderately
common.

Tylonycteris robustula Thomas, 1915.
Ann. Mag. Nat. Hist., ser. 8, 15:227.

Common Name — Greater flat-headed bat.
Greater bamboo bat.

Distribution — Southern China to the Lesser
Sunda Islands and Philippines; recorded from Lu-
zon (Rizal [usnm] and Zambales [upd] provinces),
Calauit (ummz), and Palawan (usnm).

Habitat — Disturbed lowland regions with
bamboo stands (Heaney & Alcala. 1986).

Status — Widespread, status unknown in Phil-
ippines.

Comment — First reported from the Philippines
by Heaney and Alcala (1986).

Molossidae — Free-tailed Bats

None of the four species of this family in the
Philippines (Fig. 5E) is endemic; all are poorly
known.

Chaerephon plicata (Buchanan, 1800).
Trans. Linn. Soc. Lond. 5:261.

Common Name — Wrinkle-lipped bat.

Distribution — India to Bali and Hainan. Phil-
ippine records are from Cebu (ummz), Leyte
(rom), Luzon (Cagayan [usnm], Isabela [fmnh],
Pampanga [usnm], and Rizal [fmnh] provinces),
and Negros (rom). Also reported from Mindanao
(Cotabato) by Taylor (1934).

Habitat — Recorded only in caves from sea
level to about 200 m, probably requires forested
habitat.

Status — Probably vulnerable and declining.
Formerly among the most abundant bats in some
large caves (Lawrence, 1939; Taylor, 1934); some
previously recorded colonies in caves are now de-
stroyed (Rickart et al., 1993). Reported since 1980
only from northern Luzon (Danielsen et al.,
1994).

Comment — Includes C. luzonus (Hill, 1961).

Cheiromeles parvidens Miller and Hol-
lister, 1921. Proc. Biol. Soc. Washington,
34:100.

Common Name — Lesser naked bat.

Distribution — Sulawesi and the Philippines;
reported from Mindanao (Davao del Sur [uplb],
Misamis Oriental [uplb], and South Cotabato
[amnh] provinces), and Negros (rom). Also re-
ported from Mindoro by Lawrence (1939).

Habitat — Poorly known, but probably in ag-
ricultural areas from sea level to 200 m. Often
roost in hollow trees (Rubor, 1986; Taylor. 1934).

Status — Unknown.

Comment — We recognize this as a morpholog-
ically distinctive species (Ingle & Heaney. 1992).

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

29

in contrast to Koopman (1993), who considers it
a synonym of C. torquatus.

Cheiromeles torquatus Horsfield, 1824.
Zool. Res. Java, part 8.

Common Name — Greater naked bat.

Distribution — Sumatra to Java; within the
Philippines, restricted to Palawan (fmnh).

Habitat — Unknown in the Philippines. On
Borneo, reported to roost in large caves and hol-
low trees and to feed both in open areas over
streams and clearings and above forest canopy
(Payne et al., 1985).

Status — Unknown.

Mops sarasinorum (Meyer, 1899). Abh.
Zool. Anthrop. -Ethnology, Mus. Dres-
den, 7(7): 15.

Common Name — Sulawesi mastiff bat.

Distribution — Sulawesi and the Philippines;
records are from Luzon (Benguet [fmnh]), Min-
danao (South Cotabato [amnh] and Misamis Ori-
ental [su] provinces), and Palawan (smf) only.

Habitat — Poorly known; probably found in
lowland forest.

Status — Unknown.

Comment — Freeman (1981) listed Philippinop-
terus as a synonym of Mops, and P. lanei as a
subspecies of Mops sarasinorum.

Primates

Loridae — Lorises and Coucangs

A single species of this family barely enters the
Philippines.

Nycticebus coucang Boddaert, 1785.
Elench. Anim., p. 67.

Common Name — Slow loris.

Distribution — India to Borneo and the Philip-
pines; records are from the Sulu Archipelago only
(Musser & Heaney, 1985; Timm & Birney, 1980).
Specimen from Mindanao (Misamis Occidental
[usnm]) probably is an error (Fooden, 1991a). Re-
liable records from Bongao (uplb), Sanga-sanga,

Simunul, Tawi-tawi (see Timm & Birney, 1992;
Fooden, 1991a).

Habitat — Primary and secondary lowland for-
est, gardens, and plantations (Timm & Birney,
1992; Payne et al., 1985).

Status — Widespread in Asia, but very limited
in the Philippines (Dagosto & Gebo, 1995).
CITES: Appendix II.

Tarsiidae — Tarsiers

A single species in this family occurs in the
Philippines.

Tarsius syrichta (Linnaeus, 1758). Syst.
Nat., 10th ed., 1:29.

Common Name — Philippine tarsier.

Distribution — Philippines only. Records are
from Bohol (usnm), Dinagat (dmnh), Leyte
(usnm), Mindanao (Davao del Norte [fmnh], Da-
vao del Sur [fmnh], Misamis Occidental [usnm],
Misamis Oriental [msu], South Cotabato [fmnh],
Zamboanga del Norte [fmnh], and Zamboanga del
Sur [usnm] provinces), and Samar (fmnh). Also
reported from Basilan (Lawrence, 1939), Biliran,
Maripipi (Rickart et al., 1993), and Mindanao
(Bukidnon) (Sanborn, 1953).

Habitat — Second growth, secondary forest,
and primary forest from sea level to 700 m
(Hoogstraal, 1951; Rabor, 1986; Rickart et al.,
1993; Thomas, 1898).

Status — Locally common and widespread,
largely because of its tolerance of second growth
habitat (Dagosto & Gebo, 1995). CITES: Appen-
dix II.

Comment — Reviewed by Musser and Dagosto
(1987). Physiological studies have been conduct-
ed by McNab and Wright (1987).

Cercopithecidae — Monkeys

A single species of monkey occurs in the Phil-
ippines, but is quite widespread (Fooden, 1991b).

Macaca fascicularis (Raffles, 1821).
Trans. Linn. Soc. Lond. 13:246.

Common Name — Long-tailed macaque.
Distribution — Burma to Timor; throughout the

30

FIELDIANA: ZOOLOGY

Philippines (Fooden, 1991b). Specimens from
Balabac (usnm), Basilan (usnm), Baslut (usnm),
Biliran (usnm), Bohol (fmnh), Busuanga (fmnh),
Cagayan Sulu (usnm), Culion (fmnh), Jolo
(usnm), Leyte (usnm), Luzon (Abra [fmnh], Ba-
tangas [usnm], Cagayan [usnm], Camarines Sur
[fmnh], Ilocos Norte [usnm], Isabela [amnh], and
Laguna [amnh]), Maripipi (usnm), Mindanao (Da-
vao del Norte [fmnh], Davao del Sur [fmnh], Lan-
ao del Norte [usnm], Maguindanao [fmnh], Mis-
amis Occidental [fmnh], Zamboanga del Norte
[fmnh], and Zamboanga del Sur [fmnh] prov-
inces), Mindoro (usnm), Negros (fmnh), Palawan
(fmnh), Samar (fmnh), and Sibuyan (fmnh). Also
reported from Culion, Mindanao (Davao Oriental
and Maguindanao) (Sanborn, 1952; Lawrence,
1939), Mindanao (Agusan del Norte) (Sanborn,
1953), Catanduanes (Heaney et al., 1991), Cami-
guin (Heaney & Tabarazan, 1995), Panay (Lasti-
mosa, pers. comm.), and Aurora, Quezon, and
Zambales provinces, Luzon (Ong, pers. obs.).

Habitat — Agricultural areas near forest, sec-
ond growth, secondary forest, and primary forest
from sea level to at least 1800 m in lowland and
montane forest (Dagosto & Gebo, 1995; Daniel-
sen et al., 1994; Fooden, 1991b, 1995; Goodman
& Ingle, 1993; Heaney et al., 1991, in press; Ra-
bor, 1986; Rickart et al., 1993; Thomas, 1898).

Status — Widespread in Asia. In the Philip-
pines, locally common to uncommon and hunted
heavily. CITES: Appendix II.

Comment — Geographic variation and system-
atics have been reviewed by Fooden (1991b,
1995).

Pholidota

Manidae — Pangolins

A single species of this distinctive family is
known from the Philippines.

Manis javanica Desmarest, 1822. Mam-
malogie, In Encycl. Meth. 2:377.

Common Name — Malayan pangolin.

Distribution — Burma to Java, and the Philip-
pines. Philippine records from Palawan Faunal
Region only, Palawan (fmnh) and Culion (Elera,
1915).

Habitat — Primary and secondary lowland for-

est, sometimes localized in distribution (Hoogs-
traal, 1951).

Status — Uncommon, heavily hunted and prob-
ably seriously endangered in the Philippines.

Rodentia

Sciuridae — Squirrels

All nine species of squirrels from the Philip-
pines are endemic; six species are confined to the
Palawan Faunal Region, and the remaining three
to the Mindanao Faunal Region. A representative
species is shown in Figure 4C. Hylopetes minda-
nensis is considered to be a synonym of Petino-
mys crinitus (Hoffmann et al., 1993), and Sun-
dasciurus davensis, S. mindanensis, and S. sa-
marensis are considered to be synonyms of S. phi-
lippinensis (Corbet & Hill, 1992).

Exilisciurus concinnus (Thomas, 1888).
Ann. Mag. Nat. Hist., ser. 6, 6:407.

Common Name — Philippine pygmy squirrel.

Distribution — Endemic to the Mindanao Fau-
nal Region. Recorded from the islands of Basilan
(fmnh), Biliran (usnm), Bohol (fmnh), Dinagat
(usnm), Leyte (ummz), Mindanao (Agusan del
Norte [uplb], Bukidnon [fmnh], Davao del Sur
[fmnh], Davao Oriental [dmnh], Lanao del Norte
[dmnh], Lanao del Sur [dmnh], Maguindanao
[fmnh], Misamis Occidental [fmnh]. South Cota-
bato [fmnh], Surigao del Norte [dmnh], Zamboan-
ga del Norte [fmnh], and Zamboanga del Sur
[dmnh] provinces), Samar (fmnh), and Siargao
(dmnh).

Habitat — From sea level to 2000 m, in low-
land and montane primary and secondary forest,
probably most abundant in small clearings at mid-
dle elevations (Heaney, 1985a; Rabor, 1986; Rick-
art et al., 1993).

Status — Widespread, moderately common.

Comment — Revised by Heaney (1985a).

Hylopetes nigripes (Thomas, 1893). Ann.
Mag. Nat. Hist., ser. 6, 12:30.

Common Name — Palawan flying squirrel.
Distribution — Palawan Faunal Region only.

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

BIOLOGY LIBRARY
101 BliftHiLL HAIL

31

^ -. -irtnn

Recorded on Bancalan (fmnh) and Palawan Island
(fmnh, usnm).

Habitat — Primary and secondary lowland for-
est, where they nest in cavities in large trees (Tay-
lor, 1934).

Status — Moderately common.

Petinomys crinitus Hollister, 1911. Proc.
Biol. Soc. Washington, 24:185.

Common Name — Mindanao flying squirrel.

Distribution — Mindanao Faunal Region only.
There are records from Basilan (usnm), Dinagat
(dmnh), Mindanao (Bukidnon [fmnh], Davao del
Sur [dmnh], Lanao del Sur [dmnh], Misamis Oc-
cidental [fmnh], Misamis Oriental [dmnh], and
Zamboanga del Norte [uplb] provinces), and Siar-
gao (dmnh).

Habitat — Primary lowland forest from 500 m
to 1600 m, most abundant at middle elevations
(Musser and Heaney, 1992).

Status — Moderately common in midelevation
primary forest.

Comment — Includes Hylopetes mindanensis
(Corbet & Hill, 1992; Hoffmann et al., 1993).

Sundasciurus hoogstraali (Sanborn,
1952). Fieldiana Zool., 33:115.

Common Name — Busuanga tree squirrel.

Distribution — Palawan Faunal Region. Re-
corded only from the islands of Busuanga (fmnh)
and Calauit (ummz).

Habitat — Primary and secondary lowland for-
est (Hoogstraal, 1951).

Status — Locally common.

Sundasciurus juvencus (Thomas, 1908).
Ann. Mag. Nat. Hist., ser. 8, 2:498.

Common Name — Northern Palawan tree squir-
rel.

Distribution — Palawan Faunal Region. Re-
corded from central (north of Abo-abo) and north-
ern Palawan Island only.

Habitat — Primary and secondary lowland for-
est (Hoogstraal, 1951).

Status — Locally common, stable. IUCN: En-
dangered (but we recommend delisting).

Sundasciurus moellendorffi (Matschie,
1898). Sitzb. Gesell. Naturf. Fr., Berlin,
5:41.

Common Name — Culion tree squirrel.

Distribution — Palawan Faunal Region only.
Records are from Culion (fmnh), Linapacan
(pnm), Hoc (pnm), and Tampel (pnm) islands only.

Habitat — In primary and secondary lowland
forest and coconut groves (Hoogstraal, 1951; San-
born, 1952).

Status — Locally abundant.

Comment — Currently includes S. albicauda
Matschie 1898, which is reported from Culion Is-
land, but further study is needed (Heaney, 1979;
Gonzales, unpubl. data).

Sundasciurus philippinensis (Water-
house, 1839). Proc. Zool. Soc. Lond., p.
117.

Common Name — Philippine tree squirrel.

Distribution — Philippines only, where it is
found on Mindanao and adjacent islands. Record-
ed from Basilan (mcz), Biliran (usnm), Bohol
(fmnh), Dinagat (usnm), Leyte (fmnh), Mindanao
(Agusan del Norte [uplb], Davao del Norte
[fmnh], Davao del Sur [fmnh], Davao Oriental
[uplb], Lanao del Norte [dmnh], Lanao del Sur
[dmnh], Maguindanao [fmnh], Misamis Occiden-
tal [fmnh], Misamis Oriental [dmnh], South Cot-
abato [uplb], Surigao del Sur [uplb], Zamboanga
del Norte [fmnh], and Zamboanga del Sur [dmnh]
provinces), Samar (fmnh), and Siargao (dmnh).

Habitat — In primary and secondary lowland
and montane forest from near sea level to at least
2100 m, often most abundant near agricultural
fields (Rickart et al., 1993; Sanborn, 1952).

Status — Locally common in forested regions.

Comment — Corbet and Hill (1992) synony-
mized Sundasciurus davensis, S. mindanensis, and
S. samarensis with S. philippinensis. IUCN: List-
ed S. samarensis as vulnerable, but recent taxo-
nomic changes make this invalid.

Sundasciurus rabori Heaney, 1979. Proc.
Biol. Soc. Washington, 92:281.

Common Name — Palawan montane tree squir-

rel.

Distribution — Palawan Island only (pnm).

32

FIELDIANA: ZOOLOGY

Fig. 6. Heads of representative murid rodents from Luzon (not to the same scale). A, Archboldomys luzonensis.
B, Phloeomys cumingi. C, Rhynchomys isarogensis. D, Rattus everetii.

Habitat — Recorded in mountains on Palawan
from 110 m to 1300 m.

Status — Moderately common but geographi-
cally restricted and probably confined to upper el-
evation forest. IUCN: Vulnerable.

(fmnh), Matangule (fmnh), and southern Palawan
(Brooke's Point Municipality; FMNH) islands only.

Habitat — Lowland forest, coconut groves, and
banana plantations.

Status — Common and stable.

Sundasciurus steerii (Giinther, 1877).
Proc. Zool. Soc. Lond. [1876] p. 735.

Common Name — Southern Palawan tree squir-
rel.

Distribution — Palawan Faunal Region only.
Records are from Balabac (usnm), Bancalan

Muridae — Mice and Rats

Philippine murids are a remarkably diverse
group of animals, ranging from small, ground-liv-
ing shrew-like animals to large arboreal animals
with flowing black hair (some examples are
shown in Figs. 4D and 6). An astounding 14 new

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

33

species of Philippine murid rodents have been dis-
covered since the last checklist was published in
1987. The genus Apomys includes the most new
species, with two undescribed species from Si-
buyan (Goodman & Ingle, 1993), one undescribed
species from Negros (Heaney, unpubl. data), one
from Mindoro (Dans et al., unpubl. data), and one
from Camiguin (Heaney & Tabaranza, unpubl.
data), as well as the recently described Apomys
gracilirostris from Mindoro (Ruedas, 1995). The
other additions are an undescribed Archboldomys
from northern Luzon (Rickart et al., 1998), an un-
described Bullimus from Camiguin (Heaney & Ta-
baranza, 1997), Chrotomys gonzalesi from Mt. Is-
arog, an undescribed Chrotomys from Sibuyan,
Crateromys heaneyi from Panay (Gonzales &
Kennedy, 1996), undescribed Crunomys (Rickart
et al., 1998) and Tarsomys from Mt. Kitanglad
(Heaney et al., unpubl. data), Tarsomys echinatus
from Mindanao (Musser & Heaney, 1992), and an
undescribed Tarsomys from Sibuyan (Goodman
& Ingle, 1993). In addition, two undescribed spe-
cies previously known (Batomys from Dinagat
[Musser et al., in press] and Haeromys from Pa-
lawan [Heaney et al., 1987]) are curently under
study (Musser et al., unpubl. data).

The systematics and relationships of Philippine
murid genera were reviewed by Musser and Hea-
ney (1992). Chromosomal variation in Philippine
murids was described by Rickart and Musser
(1993). The species formerly referred to as Rattus
rattus is now recognized as a species group, and
Philippine populations have been placed in the
species Rattus tanezumi (Musser & Carleton,
1993). Bullimus rabori and Rattus tyrannus are
now considered to be junior synonyms of Bulli-
mus bagobus and Rattus everetti, respectively
(Musser & Carleton, 1993; Musser & Heaney,
1992). Crunomys rabori is now considered to be
a synonym of C. melanius (Musser & Heaney,
1992; Rickart et al., unpubl. data).

Abditomys latidens (Sanborn, 1952).
Fieldiana Zool., 33:125.

Common Name — Luzon broad-toothed rat.

Distribution — Luzon Faunal Region. Appar-
ently endemic to central and northern Luzon Is-
land (Laguna [usnm] and Mountain [fmnh] prov-
inces).

Habitat — Very poorly known. Taken in dense-
ly vegetated gullies in pine forest at about 2250
m (Rabor, 1955, 1986; Sanborn, 1952) and in sec-

ond growth in the lowlands at about 75 m (Bar-
behenn et al., 1973; Musser, 1982a).
Status — Uncertain.

Anonymomys mindorensis Musser, 1981.
Bull. Am. Mus. Nat. Hist., 168:300.

Common Name — Mindoro climbing rat.

Distribution — Endemic to Mindoro (fmnh)
and known only from Hong Peak, Halcon Range.

Habitat — Poorly known; taken in forest at
1400 m (Musser, 1981).

Status — Uncertain; geographically restricted
to an island that has been heavily deforested. It
may be vulnerable to further deforestation (Hea-
ney & Utzurrum, 1991). IUCN: Vulnerable.

Apomys abrae (Sanborn, 1952). Fieldi-
ana Zool., 33(2): 133.

Common Name — Luzon Cordillera forest
mouse.

Distribution — Endemic to the Central Cordil-
lera of northern Luzon (Abra [fmnh], Benguet
[fmnh], Ilocos Norte [fmnh], and Mountain
[fmnh] provinces).

Habitat — Moderately common in primary for-
est and second growth from ca. 1000 m to 2000
m, occasionally to 2500 m (Musser, 1982c; Rabor,
1955; Sanborn, 1952).

Status — Probably stable.

Comment — The genus Apomys was reviewed
by Musser (1982c), with modifications by Musser
and Heaney (1992).

Apomys datae (Meyer, 1899). Abh. Mus.
Dresden, ser. 7, 7:25.

Common Name — Luzon montane forest mouse.

Distribution — Endemic to northern Luzon
(Benguet [usnm], Cagayan [fmnh], Ilocos Norte
[su], Isabela [pnm], and Mountain [fmnh] prov-
inces).

Habitat — Recorded in primary montane and
mossy forest from 760-m to 1650-m elevation in
the Sierra Madre (Danielsen et al., 1994; Mallari
& Jensen, 1993) and from 1600 m to 2500 m in
montane and mossy forest in the Central Cordil-
lera (Musser, 1982b; Rabor, 1955; Sanborn,
1952).

Status — Moderately common and widespread.

34

FIELDIANA: ZOOLOGY

Apomys gracilirostris Ruedas, 1995.
Proc. Biol. Soc. Washington, 108:305.

Apomys littoralis (Sanborn, 1952). Field-
iana Zool., 33(2): 134.

Common Name — Large Mindoro forest mouse.

Distribution — Known only from Mindoro
(Ruedas, 1995).

Habitat — Recorded from 1255 m to 1900 m in
primary montane forest and natural bamboo thick-
et (Ruedas, 1995).

Status — Locally common, but seriously threat-
ened by extensive habitat loss (Ruedas, 1995).
IUCN: Vulnerable.

Apomys hylocoetes Mearns, 1905. Proc.
U.S. Natl. Mus., 28:456.

Common Name — Mindanao mossy forest
mouse.

Distribution — Endemic to Mindanao (Bukid-
non [fmnh] and Davao del Sur [usnm] provinces).

Habitat — Uncommon in primary montane for-
est at 1900 m and abundant in primary mossy
forest from 2250 m to 2800 m (Musser, 1982c;
Heaney et al., unpubl. data). Probably widespread
on the high peaks of Mindanao.

Status — Stable and probably moderately wide-
spread.

Apomys ins ignis Mearns, 1905. Proc.
U.S. Natl. Mus., 28:459.

Common Name — Mindanao montane forest
mouse.

Distribution — Endemic to the Mindanao Fau-
nal Region: Dinagat (dmnh) and Mindanao (Bu-
kidnon [fmnh), Davao del Norte [fmnh], Davao
del Sur [fmnh), Misamis Oriental [fmnh], Mis-
amis Occidental [fmnh], and Zamboanga del Nor-
te [amnh] provinces). Also reported from Min-
danao (Agusan del Norte) (Sanborn, 1953).

Habitat — In primary and secondary forest
from 900 m to 2800 m (Musser & Heaney, 1992).
On Mt. Kitanglad, absent from lowland forest
(1100 m), uncommon in lower montane forest
(1600 m), abundant in upper montane forest (1800
m) and transitional montane-mossy forest (1900
m), and absent in mossy forest (2250-2800 m)
(Heaney et al., unpubl. data).

Status — Stable and widespread.

Common Name — Mindanao lowland forest
mouse.

Distribution — Endemic to the Mindanao Fau-
nal Region: Biliran (usnm), Bohol (usnm), Leyte
(usnm), and Mindanao (Lanao del Sur [hmnh] and
Maguindanao [fmnh] provinces).

Habitat — The holotype was taken on the
coastal plain of northern Cotabato (now Maguin-
danao) Province. An additional specimen taken at
about 1400 m in Lanao del Sur was tentatively
referred to this species (Musser, 1982c). Mice ten-
tatively referred to this species were common on
Leyte in primary montane and mossy forest and
on Biliran in montane forest and were rare in low-
land forest on Leyte and Bohol (Heaney et al.,
1989; Rickart et al., 1993; Heaney et al., unpubl.
data). Specimens initially referred to this species
from Negros (Musser, 1982c) are an undescribed
species (Musser & Heaney, 1992) listed here as
Apomys sp. A.

Status — Uncertain because of taxonomic un-
certainty.

Apomys microdon Hollister, 1913. Proc.
U.S. Natl. Mus., 46:327.

Common Name — Small Luzon forest mouse.

Distribution — Endemic to the Luzon Faunal
Region: Catanduanes (usnm) and Luzon (Cama-
rines Sur [usnm] and Isabela [fmnh] provinces).

Habitat — Uncommon on Mt. Isarog. where
they were captured in secondary lowland forest
and primary montane forest from 475 m to 1550
m (Heaney et al., in press; Rickart et al., 1991).
They were also taken in forest near sea level in
Isabela Province (Danielsen et al., 1994) and on
Catanduanes (Hollister, 1913). The species is vir-
tually unknown elsewhere (Heaney et al.. 1991;
Mallari & Jensen, 1993).

Status — Uncertain; probably widespread and
stable, but deforestation has removed much of its
habitat.

Comment— Corbet and Hill (1992) stated that
A. hollisteri should be used for this species, but
Musser and Carleton (1993) continued usage of
A. microdon.

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

35

Apomys musculus Miller,
U.S. Natl. Mus., 38:403.

1911. Proc. Apomys sp. B

Common Name — Least Philippine forest
mouse.

Distribution — Endemic to Dinagat (fmnh) and
Luzon (Benguet [usnm], Camarines Sur [fmnh],
and Isabela [fmnh] provinces), and Mindoro
(fmnh).

Habitat — Abundant in primary montane forest
(1125-1350 m), uncommon in mossy forest
(1550-1750 m), and absent in disturbed lowland
forest (475-900 m) on Mt. Isarog, southern Luzon
(Balete & Heaney, in press; Heaney et al., in
press; Rickart et al., 1991). Recorded in disturbed
lowland forest at 300 m in northeastern Luzon
(Danielsen et al., 1994; Mallari & Jensen, 1993)
and at 1500 m in Benguet (Miller, 1910) and in
ridgetop mossy forest at 750 m on Dinagat (Ta-
baranza, unpubl. data).

Status — Apparently widespread and moderate-
ly common at medium elevations.

Common Name — Greater Sibuyan forest
mouse.

Distribution — Sibuyan only (fmnh; Goodman
& Ingle, 1993).

Habitat — Primary forest from 325 m to 1325
m (Goodman & Ingle, 1993; specimens in fmnh).

Status — Locally common, but habitat is lim-
ited and declining (Goodman & Ingle, 1993).

Apomys sp. C

Common Name — Lesser Sibuyan forest mouse.

Distribution — Sibuyan only (Goodman & In-
gle, 1993).

Habitat — Primary forest from 30 m to 325 m
(Goodman & Ingle, 1993; specimens in fmnh).

Status — Locally abundant, but habitat is very
limited and declining (Goodman & Ingle, 1993).

Apomys sacobianus Johnson, 1962. Proc.
Biol. Soc. Washington, 75:318.

Common Name — Long-nosed Luzon forest
mouse.

Distribution — Endemic to Luzon (Isabela
[fmnh], Mountain [usnm], Pampanga [usnm], and
Zambales [pnm] provinces).

Habitat — Poorly known, but it apparently oc-
curs in primary forest from about 200 m to 1100
m (Danielsen et al., 1994; Johnson, 1962; Mallari
& Jensen, 1993; Musser, 1982c).

Status — Uncertain. The available data indicate
that it is widespread in northern and central Lu-
zon. IUCN: Vulnerable (but we consider this list-
ing to be premature, given the current lack of
knowledge).

Apomys sp. A

Common Name — Western Visayan forest
mouse.

Distribution — Known only from Negros
(usnm) and Panay (pnm).

Habitat — Montane and mossy primary forest
from 800 m to 1600 m (Heaney et al., 1989; Hei-
deman et al., 1987; Rabor et al., 1970).

Status — Locally common in upland forest but
seriously threatened by habitat loss.

Apomys sp. D

Common Name — Camiguin forest mouse.

Distribution — Camiguin Island only (fmnh).

Habitat — Primary montane forest from 1000
m to 1275 m (Heaney & Tabaranza, 1997).

Status — Locally common and stable but con-
fined to a very small area that is threatened by
deforestation (Heaney & Tabaranza, 1997).

Apomys sp. E

Common Name — Lesser Mindoro forest mouse.
Distribution — Mindoro Island only (uplb).
Habitat — Unknown.
Status — Unknown .

Comment — Currently under study by Dans et
al. (unpubl. data).

Archboldomys luzonensis Musser, 1982.
Bull. Am. Mus. Nat. Hist., 174:30.

Common Name — Isarog shrew-mouse.

Distribution — Endemic to Mt. Isarog, Cama-
rines Sur Province, Luzon (fmnh).

Habitat — Moderately common in primary
montane and mossy forest from 1350 m to 1750
m (Balete & Heaney, in press; Heaney et al., in

36

FIELDIANA: ZOOLOGY

press; Rickart et al., 1991). Holotype stated to be
from approximately 750 m (Musser, 1982c).

Status — Population stable but highly restricted
in distribution, threatened by habitat destruction
(Heaney & Utzurrum, 1991). IUCN: Endangered.

Archboldomys sp. A

Common Name — Palanan shrew-mouse.

Distribution — Known only from Mt. Cetaceo.
Cagayan Province, Luzon (fmnh).

Habitat — Mossy forest at 1650 m (Danielsen
et al., 1994).

Status — Unknown.

Comment — Mistakenly identified by Heaney as
Crunomys fallax in Danielsen et al. (1994). Cur-
rently being described by Rickart et al. (1998).

Batomys dentatus Miller, 1910. Proc.
U.S. Natl. Mus., 38:400.

Common Name — Large-toothed hairy-tailed
rat.

Distribution — Known only from Benguet
Province (usnm), Luzon.

Habitat — The single known specimen was
taken at ca. 2100 m, probably in montane forest
(Miller, 1910).

Status — Unknown.

Batomys granti Thomas, 1895. Ann.
Mag. Nat. Hist., ser. 6, 16:162.

Common Name — Luzon hairy-tailed rat.

Distribution — Luzon Faunal Region; known
only from Mt. Data, Benguet Province (fmnh) and
Mt. Isarog, Camarines Sur Province (fmnh).

Habitat — Uncommon in montane forest and
common in mossy forest on Mt. Isarog (Balete &
Heaney, in press; Heaney et al., in press; Rickart
et al., 1991). Found in thick vegetation on the
ground in mossy forest at ca. 2100 m in the Cen-
tral Cordillera (Rabor, 1955; Sanborn, 1952; Tho-
mas, 1898).

Status — Probably widespread and moderately
common at higher elevations.

Batomys salomcmseni (Sanborn, 1953).
Vidensk. Medd. Nat. Foren. Kjobenhavn,
115:287.

Common Name — Mindanao hairy-tailed rat.
Distribution — Mindanao Faunal Region. Re-

cords are from Biliran (USNM), Dinagat (mm
Leyte (usnm), and Mindanao (Bukidnon [fm\h]>.

Habitat — On Leyte, common in montane and
ridgetop mossy primary forest from 7(X) m to 950
m and uncommon at 500 m in lowland forest
(Heaney et al., 1989; Rickart et al., 1993). On Mt.
Kitanglad, Mindanao, moderately common in
montane and mossy forest from 1600 m to 2375
m (Heaney et al., unpubl. data).

Status — Probably widespread and common in
primary montane and mossy forest.

Comment — Mindanaomys is considered to be a
synonym of Batomys (Musser & Heaney. 1992).

Batomys sp. A

Common Name — Dinagat hairy-tailed rat.

Distribution — Known only from Dinagat Is-
land (dmnh) (Heaney and Rabor, 1982).

Habitat — Disturbed lowland forest at ca. 350
m (Musser et al., unpubl. data).

Status — Unknown; geographically restricted
and habitat subject to destruction.

Comment — Currently being described by Mus-
ser et al. (in press).

Bullimus bagobus Mearns, 1905. Proc.
U.S. Natl. Mus., 28:450.

Common Name — Large Mindanao forest rat.

Distribution — Widespread in Mindanao Fau-
nal Region; records from Bohol (fmnh), Dinagat
(dmnh), Leyte (usnm), Maripipi (usnm), Minda-
nao (Agusan del Norte [dmnh|. Davao del Sur
[fmnh], Lanao del Norte [uplbj, Lanao del Sur
[uplb], Misamis Occidental [fmnh). Misamis Ori-
ental [uplbJ, South Cotabato [amnh], Surigao del
Sur [uplb], and Zamboanga del Norte [fmnh]
provinces), Samar (usnm), and Siargao (dmnh).

Habitat — Recorded from ca. 200 m to 1800 m
on Mindanao (Musser & Heaney. 1992), mostly
in lowland forest though occasionally in mossy
forest (Sanborn, 1952), from 300 m to 500 m in
lowland forest on Leyte. and in montane forest at
740 m on Maripipi (Rickart et al.. 1993).

Status — Common and widespread.

COMMENT — The genus Bullimus was recog-
nized and defined by Musser (1982c) and Musser
and Heaney (1992). Includes Bullimus rabori
(Musser, 1982c; Musser & Carleton. 1993; Mus-
ser & Heaney. 1992).

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

37

Bullimus luzonicus (Thomas, 1895).
Ann. Mag. Nat. Hist., ser. 6, 16:163.

Common Name — Large Luzon forest rat.

Distribution — Known only from Luzon (Au-
rora [upd], Benguet (fmnh), and Camarines Sur
[fsm] provinces).

Habitat — Poorly known. Scattered specimens
taken in mixed primary and distubed lowland for-
est from about 200 m to 2400 m in montane and
mossy forest (Largen, 1985; Musser, 1982c; Ra-
bor, 1955; Sanborn, 1952; Thomas, 1898; Heaney
et al., in press).

Status — Uncertain; may be moderately com-
mon in primary lowland forest.

Bullimus sp. A

Common Name — Camiguin forest rat.

Distribution — Camiguin Island only (fmnh).

Habitat — Primary montane and mossy forest
from 1000 m to 1475 m (Heaney & Tabaranza,
1997).

Status — Stable but very restricted geographi-
cally and threatened by logging.

Comment — Initially, but incorrectly, identified
as a species of Tarsomys (Heaney & Tabaranza,
1997; Heaney et al., 1997).

Carpomys melanurus Thomas, 1895.
Ann. Mag. Nat. Hist., ser. 6, 16:162.

Common Name — Short-footed Luzon tree rat.

Distribution — Known only from Mt. Data,
Benguet Province, Luzon (fmnh).

Habitat — Montane and mossy forest from
about 2200 m to 2500 m (Largen, 1985; Thomas,
1898).

Status — Unknown; may be locally common
(Thomas, 1898), but may be limited in distribu-
tion.

Carpomys phaeurus Thomas, 1895. Ann.
Mag. Nat. Hist., ser. 6, 16:162.

Common Name — White-bellied Luzon tree rat.

Distribution — Northern Luzon only, recorded
from Benguet and Ifugao provinces (fmnh).

Habitat — Mossy forest from ca. 2200 m to
2500 m (Largen, 1985; Rabor, 1955; Sanborn,
1952; Thomas, 1898).

Status — Unknown; probably limited in distri-
bution.

Celaenomys silaceus (Thomas, 1895).
Ann. Mag. Nat. Hist., ser. 6, 16:161.

Common Name — Blazed Luzon shrew-rat.

Distribution — Northern Luzon only. Records
are from high elevation forest in Benguet (fmnh).

Habitat — In thick vegetation in mossy forest
at 2200 m to 2500 m (Sanborn, 1952; Thomas,
1898).

Status — Uncertain; may be moderately com-
mon (pnm specimens).

Comment — Included within the genus Chroto-
mys by Corbet and Hill (1992).

Chiropodomys calamianensis (Taylor,
1934). Monogr. Bur. Sci. Manila., 30:
470.

Common Name — Palawan pencil-tailed tree
mouse.

Distribution — Palawan Faunal Region only
(Musser, 1979). Records are from Balabac (usnm),
Busuanga (uimnh), and Palawan (fmnh) islands.

Habitat — Known only from lowland forest
near sea level (Taylor, 1934), coconut groves, and
bamboo thickets (Sanborn, 1952).

Status — Unknown.

Chrotomys gonzalesi Rickart and Hea-
ney, 1991. Proc. Biol. Soc. Washington,
104:389.

Common Name — Isarog striped shrew-rat.

Distribution — Known only from Mt. Isarog,
southern Luzon (fmnh).

Habitat — Occurs in montane and mossy forest,
ca. 1350 m and above, on Mt. Isarog (Balete &
Heaney, in press; Heaney et al., in press; Rickart
& Heaney, 1991; Rickart et al., 1991).

Status — Population stable but highly restricted
in distribution, threatened by habitat destruction.
IUCN: Critically endangered.

Chrotomys mindorensis Kellogg, 1945.
Proc. Biol. Soc. Washington, 58:123.

Common Name — Lowland striped shrew-rat.
Distribution — Philippines only. Recorded

38

FIELDIANA: ZOOLOGY

from Mindoro (fmnh) and lowland central Luzon
(Laguna Province [usnm]). Also reported from
Luzon (Nueva Ecija, Pampanga, and Tarlac prov-
inces) (Barbehenn et al., 1973).

Habitat — Primary and secondary forest, oc-
casionally in adjacent agricultural areas, from near
sea level to at least 1000 m (Barbehenn et al.,
1973; Kellogg. 1945; Musser et al., 1981; Rickart
& Heaney, 1991).

Status — Widespread in forest, but adversely
affected by habitat destruction.

Chrotomys whiteheadi Thomas, 1895.
Ann. Mag. Nat. Hist., ser. 6, 16:161.

Common Namk — Luzon montane striped
shrew-rat.

Distribution — Luzon only, records are from
Benguet (fmnh) and Mountain provinces (pnm).

Habitat — Recorded in mossy forest from 2300
m to 2500 m in the Central Cordillera (Largen,
1985; Rabor, 1955; Sanborn, 1952).

Status — Uncertain. It may be moderately com-
mon and widespread in the Central Cordillera.
IUCN: Vulnerable.

Chrotomys sp. A

Common Namk — Sibuyan striped shrew-rat.

Distribution — Known only from near sea level
on Sibuyan Island (fmnh).

Habitat — Lowland forest on Sibuyan (Good-
man & Ingle. 1993; specimens in fmnh).

Status — May be restricted to a small patch of
lowland forest on Sibuyan (Goodman & Ingle,
1993).

Crateromxs australis Musser, Heaney, and
Rabor, 1985. Am. Mus. Novit., 2821:3.

Common Namf — Dinagat hairy-tailed cloud
rat.

Habitat — Known only from a single specimen
from Dinagat Island (Musser et al., 1985; Oliver
et al., 1993a).

Habitat — Poorly known; apparently from low-
land forest.

Status — Unknown; geographically restricted.
This species is probably badly affected by defor-
estation (Oliver et al., 1993a). IUCN: Endangered.

Crateromys heanevi Gonzales and Ken-
nedy, 1996. J. Mammal., 76:26.

Common Namk — Panay bushy-tailed cloud rat,
Panay cloud runner.

Distribution — Panay only (Gonzales & Ken-
nedy, 1996; Oliver et al., 1993a).

Habitat — Lowland primary and secondary for-
est to about 400 m (Gonzales & Kennedy, 1996;
Oliver et al., 1993a).

Status — Severely impacted by habitat destruc-
tion on Panay. Highly endangered (Gonzales &
Kennedy, 1996; Oliver et al.. 1993a). IUCN: En-
dangered.

Crateromys paulus Musser and Gordon,
1981. J. Mammal., 62:515.

Common Namk — Ilin hairy-tailed cloud rat.

Distribution — Known with certainty only from
Ilin Island (south of Mindoro; usnm), but unveri-
fied reports suggest its presence on southern Min-
doro (Oliver et al., 1993a).

Habitat — Unknown; probably lowland forest
(Musser & Gordon, 1981).

Status — Uncertain; geographically extremely
restricted (Heaney & Utzurrum, 1991). Reported
extinct on Ilin (Pritchard, 1989). Unverified re-
ports indicate its possible occurrence on Mindoro
(Oliver, 1994; Oliver et al., 1993a). IUCN: Criti-
cally endangered.

Crateromys schadenbergi (Meyer, 1895).
Abh. Mus. Dresden, 6:1.

Common Namk — Luzon bushy-tailed cloud rat.

Distribution — Known only from Benguet. Ifu-
gao, and Mountain provinces, Luzon (fmnh; Oli-
ver et al., 1993a; Sanborn, 1952).

Habitat — From 2000 m to 2500 m in pine and
mossy forest in the central Cordillera (Rabor.
1955; Sanborn, 1952).

Status — Apparently locally common in oak-
pine forest, rare elsewhere; hunted. IUCN: Vul-
nerable.

Crunomxs fallax Thomas, 1897. Trans.
Zool. Soc. Lond. 14(6):394.

Common Nami — Northern Luzon shrew-
mouse.

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

39

Distribution — Known only from a single spec-
imen taken at about 300 m in the Sierra Madre of
northern Luzon (Isabela Prov. [bmnh]).

Habitat — Unknown; possibly confined to pri-
mary lowland forest (Musser, 1982c; Thomas,
1898).

Status — Unknown; probably dependent on
lowland forest, which has diminished greatly on
Luzon. IUCN: Critically endangered. We believe
this listing to be premature, given the current lack
of knowledge.

Comment — A specimen from Cagayan Prov-
ince, Luzon erroneously identified by Heaney as
this species (Danielsen et al., 1994) is now being
described as a new species of Archboldomys
(Rickart et al., 1998).

Crunomys melanius Thomas, 1907. Proc.
Zool. Soc. Lond., p. 141.

Common Name — Southern Philippine shrew-
mouse.

Distribution — Known only from Camiguin
(Heaney & Tabaranza, 1995; fmnh), Leyte
(dmnh), and Mindanao (Bukidnon [fmnh], Cota-
bato [uplb], and Davao del Norte [bmnh] prov-
inces).

Habitat — From near sea level to 900 m, prob-
ably in primary rain forest (Musser & Heaney,
1992; Heaney & Tabaranza, 1995).

Status — Widespread and stable in good habi-
tat, but restricted to lowland forest, which has
been greatly reduced.

Comment — Includes the Leyte shrew-mouse
{Crunomys rabori Musser, 1982) (Musser & Hea-
ney, 1992; Rickart et al., 1998).

Crunomys sp. A

Common Name — Kitanglad shrew-mouse.

Distribution — Known only from a single spec-
imen taken at 2250 m on Mt. Kitanglad, Minda-
nao (Bukidnon Province [fmnh]).

Habitat — Primary mossy forest (Rickart et al.,
1998).

Status — Unknown.

Haeromys sp. A

Common Name — Palawan pygmy tree mouse.
Distribution — Known only from Palawan and

Calauit islands in the Palawan Faunal Region
(Musser & Newcomb, 1983).

Habitat — Unknown.

Status — Unknown.

Limnomys sibuanus Mearns, 1905. Proc.
U.S. Natl. Mus., 28:452.

Common Name — Long-tailed moss-mouse.

Distribution — Endemic to Mindanao Island.
Recorded from Bukidnon (fmnh), Davao del Sur
(usnm), and Misamis Occidental (usnm) prov-
inces.

Habitat — From 2000 m to 2800 m in primary
montane and mossy rainforest (Musser, 1994;
Musser & Heaney, 1992).

Status — Common in high-elevation forest
(Heaney et al., unpubl. data).

Comment — The genus Limnomys was reviewed
and redefined by Musser ( 1 977b) and Musser and
Heaney (1992).

Maxomys pang lima (Robinson, 1921).
Ann. Mag. Nat. Hist., ser. 9, 7:235.

Common Name — Palawan spiny rat.

Distribution — Palawan Faunal Region only.
Recorded from Balabac (usnm), Busuanga
(fmnh), Calauit (uplb), Culion (fmnh), and Pala-
wan (fmnh).

Habitat — Secondary and primary forest from
near sea level to at least 1000 m (Barbehenn et
al., 1973; Hoogstraal, 1951; Musser et al., 1979;
Sanborn, 1952).

Status — Common .

Comment — The genus Maxomys was reviewed
by Musser et al. (1979) and in part by Ruedas and
Kirsch (1997).

Mus musculus Linnaeus, 1758. Syst.
Nat, 10th ed., 1:62.

Common Name — House mouse.

Distribution — Nearly worldwide; widespread
in Southeast Asia. Found throughout the Philip-
pines; specimens from Bohol (usnm), Leyte
(dmnh), Luzon (Laguna [uplb] Province), Mari-
pipi (usnm), and Negros (usnm). Also reported
from Mindanao (Davao del Norte Province) by
Barbehenn et al. (1973).

Habitat — Human habitations in urban and ru-

40

FIELDIANA: ZOOLOGY

ral areas, rarely above 100 m elevation (Heaney
et al., in press; Rabor, 1986).

Status — Non-native and abundant.

Comment — All Philippine populations of Mus
are now placed in the species M. musculus (sub-
species castaneus) and the species is considered
to be non-native (Marshall, 1977, 1986; Marshall
& Sage, 1981).

Palawanomys furvus Musser and New-
comb, 1983. Bull. Am. Mus. Nat. Hist.,
174:335.

Common Name — Palawan soft-furred mountain
rat.

Distribution — Palawan Faunal Region; known
only from Mt. Mantalingajan, Palawan Island
(usnm).

Habitat — Unknown, but probably high moun-
tain forest (Musser & Newcomb, 1983).

Status — Poorly known; probably highly re-
stricted in distribution, possibly threatened by log-
ging. IUCN: Endangered.

Phloeomys cumingi (Waterhouse, 1839).
Proc. Zool. Soc. Lond. p. 108.

Common Name — Bugkun, Southern Luzon gi-
ant cloud rat.

Distribution — Luzon Faunal Region only. Re-
corded on Catanduanes (usnm) and southern Lu-
zon (Camarines Sur [fmnh], Laguna [uplb], and
Quezon [uplb] provinces; Oliver et al., 1993a).
Reported from Marinduque (Steere, 1890).

Habitat — On Mt. Isarog and Catanduanes, this
species was found in disturbed lowland forest
from 150 m to 900 m elevation (Heaney et al.,
1991, Heaney et al., in press; Oliver et al.. 1993a;
Rickart et al., 1991).

Status — Moderately widespread and common,
but heavily hunted and subject to habitat destruc-
tion (Oliver et al., 1993a). IUCN: Vulnerable.

Phloeomys pallidus Nehring, 1890.
I Sitzb. Ges. Naturf. Fr. Berlin, p. 106.

Common Name — Bu-ot, Northern Luzon giant
cloud rat.

Distribution — Widespread in northern and
central Luzon (Oliver et al., 1993a). Verified re-
cords from Abra (fmnh), Benguet (usnm), Kalin-

ga-Apayao (amnh), Laguna (UPLB), and Nueva
Viscaya (uplb) provinces (Oliver et al., 1993a).
Recently verified from Bataan/Zambales region
(Ong, unpubl. data).

Habitat — From sea level to high mountains (at
least 2000 m), in primary and secondary forest
(Rabor, 1955; Thomas, 1898) and heavily dis-
turbed scrub (Oliver et al.. 1993a).

Status — Widespread and apparently common
in forests; hunted (Oliver et al.. 1993a; Pasicolan,
1993).

Rattus argentiventer (Robinson & Kloss,
1916). J. Strs. Br. Roy. Asiat. Soc., 73:
274.

Common Name — Rice-field rat.

Distribution — Thailand to New Guinea. In the
Philippines, recorded on Cebu (uplb), Luzon (La-
guna [uplb] Province), Mindanao (Davao del Sur
[fmnh] Province), Mindoro (fmnh), and Negros
(uplb).

Habitat — Rice-fields, grasslands, and planta-
tions (Payne et al., 1985).

Status — Non-native, may be locally abundant.

Comment — Originally reported from the Phil-
ippines as R. rattus umbriventer Kellogg 1945
(Barbehenn et al., 1973; Musser, 1973).

Rattus everetti (Giinther, 1879). Proc.
Zool. Soc. Lond. p. 75.

Common Name — Common Philippine forest
rat.

Distribution — Endemic but widespread in the
Philippines, excluding the Palawan and Sulu fau-
nal regions and the Batanes/Babuyan groups.
Specimens are recorded from Biliran (usnm), Bo-
hol (fmnh), Camiguin (DMNH), Catanduanes
(fmnh), Dinagat (usnm), Leyte (usnm). Luzon
(Abra [fmnh], Albay [uplb], Aurora [updJ, Ben-
guet [fmnh], Camarines Sur [fmnh]. Laguna
[usnm], Mountain Province [fmnh], Pampanga
[amnh], Quezon [uplb], Rizal [uplb], and Sorso-
gon [fmnh] provinces). Marinduque (upd). Mari-
pipi (usnm). Mindanao (Bukidnon [fmnh], Davao
del Norte [fmnh], Davao del Sur [fmnh], Lanao
del Norte [uplb], Lanao del Sur [uplb], Maguin-
danao [fmnh]. Misamis Occidental [fmnh]. Mis-
amis Oriental [uplb]. South Cotabato [amnh], Su-
rigao del Norte [uplb], Surigao del Sur [uplb).
and Zamboanga del Norte [fmnh) provinces).

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

41

Mindoro (fmnh), Panay (pnm), Siargao (dmnh),
and Ticao (usnm). Also reported from Samar
(Johnson, 1946).

Habitat — Found in primary and disturbed low-
land, montane, and mossy forest, from sea level
to 2200 m on Luzon (Balete & Heaney, in press;
Danielsen et al., 1994; Heaney et al., 1991, in
press; Rabor, 1955) and up to 2400 m on Min-
danao (Musser & Heaney, 1992; Rickart et al.,
1993).

Status — Common in primary forest, uncom-
mon in secondary forest, and usually absent in
agricultural areas.

Comment — As currently defined, includes R.
albigularis, R. gala, R. tagulayensis, and R. tyr-
annus (Musser & Heaney, 1992; Musser & Carle-
ton, 1993).

Rattus exulans (Peale, 1848). Mammalia.
In Repts. U.S. Expl. Surv., 8:47.

Common Name — Polynesian rat, Small spiny
rice-field rat.

Distribution — Bangladesh to Easter Island and
throughout the Philippines. Specimens taken from
Balabac (usnm), Biliran (ummz), Bohol (usnm),
Busuanga (usnm), Camiguin (dmnh), Catanduanes
(usnm), Cebu (uplb), Culion (usnm), Dinagat
(usnm), Leyte (uplb), Luzon (Laguna [usnm]
Province), Marinduque (pnm), Mindanao (Davao
del Sur [uplb], Lanao del Norte [uplb], Lanao del
Sur [uplb], Misamis Oriental [uplb], South Cot-
abato [uplb], Surigao del Norte [uplb], and Suri-
gao del Sur [uplb] provinces), Negros (usnm), and
Palawan (usnm). Also reported from Caluya Is-
land (Alcala & Alviola, 1970).

Habitat — Agricultural areas throughout the
country at all elevations (Barbehenn et al., 1973;
Rabor, 1986). Often present in disturbed forest
(e.g., Danielsen et al., 1994) and usually rare in
primary forest, but may be common in primary
forest on islands such as Negros with few native
rodents (Heaney et al., 1989).

Status — Non-native and abundant.

Rattus mindorensis (Thomas, 1898).
Trans. Zool. Soc. Lond., 14:402.

Common Name — Mindoro soft-furred rat.
Distribution — Known only from Mindoro Is-
land (fmnh).

Habitat — Forest at 1000 m to 1500 m (Tho-
mas, 1898).

Status — Apparently common at high eleva-
tions, but adversely affected by forest destruction.
IUCN: Vulnerable.

Rattus nitidus (Hodgson, 1845). Ann.
Mag. Nat. Hist., [ser. 1], 15:267.

Common Name — Himalayan field rat.

Distribution — Nepal to New Guinea. In the
Philippines, known only from Benguet Province,
Luzon (fmnh).

Habitat — Generally in houses in hilly areas
(Lekagul & McNeely, 1977).

Status — Non-native. May be locally abundant
in highland agricultural areas, but few records
from the Philippines.

Rattus norvegicus (Berkenhout, 1769).
Outlines of the Natural History of Great
Britain and Ireland, 1:5.

Common Name — Common brown rat.

Distribution — Worldwide. In the Philippines,
reported from Luzon (Bulacan, Laguna, Nueva
Ecija, and Pampanga provinces) by Barbehenn et
al. (1973).

Habitat — In the Philippines, primarily restrict-
ed to large cities and places where large ships
dock (Rabor, 1986).

Status — Non-native and abundant in urban ar-

Rattus tanezumi Temminck, 1 844. In Sie-
bold, Temminck, and Schlegel, Fauna Ja-
ponica, Arnz et Socii, Lugduni Batavo-
rum, p. 51.

Common Name — Oriental house rat.

Distribution — Afghanistan, Indo-malaya, New
Guinea, and Micronesia (except the Samoas).
Throughout the Philippines; recorded from Biliran
(usnm), Bohol (usnm), Calauit (ummz), Camiguin
(dmnh), Catanduanes (usnm), Cebu (uplb), Dina-
gat (usnm), Leyte (usnm), Luzon (Cagayan
[ummz], Camarines Sur [upd], Laguna [usnm],
Quezon [ummz], Sorsogon [fmnh], Tarlac [uplb],
and Zambales [usnm] provinces), Marinduque
(pnm), Maripipi (usnm), Mindanao (Agusan del
Norte [uplb], Bukidnon [uplb], Davao del Norte

42

FIELDIANA: ZOOLOGY

[uplb], Lanao del Norte [uplb], Lanao del Sur
[uplb], Misamis Occidental [uplbJ, Misamis Ori-
ental [uplb], North Cotabato [usnm], South Cot-
abato [uplb], Surigao del Norte [uplb], and Zam-
boanga del Norte [uplb] provinces), Mindoro
(uplb), Negros (usnm), Panay (su), Siargao
(dmnh), and Siquijor (su). Also reported from
Caluya, Sibay, Semirara, Boracay, and Carabao
islands (Alcala & Alviola, 1970).

Habitat — Abundant in urban and agricultural
areas, common in disturbed lowland and montane
forest up to 1800 m (Danielsen et al., 1994; Hea-
ney et al., 1989, in press; Heaney & Tabaranza,
1995; Rabor, 1986; Sanborn, 1952).

Status — Non-native. Abundant.

Comment — Formerly included within Rattus
rattus (Musser & Carleton, 1993). Includes many
populations formerly recognized as distinct spe-
cies (Musser, 1977a), including Rattus mindanen-
sis.

Rattus tawitawiensis Musser and Heaney,
1985. Am. Mus. Novit. 2818:5.

Common Name — Tawi-tawi forest rat.

Distribution — Known only from Tawi-tawi Is-
land, Sulu Archipelago.

Habitat — Unknown.

Status — Uncertain; geographically restricted,
probably threatened by habitat destruction. IUCN:
Vulnerable.

Rattus tiomanicus (Miller, 1900). Proc.
Washington Acad. Sci., 2:209.

Common Name — Malaysian field rat.

Distribution — Malay Peninsula to Borneo and
Palawan. In the Philippines, in Palawan Faunal
Region only. Recorded from Arena (fmnh), Ban-
calan (fmnh), Busuanga (fmnh), Calauit (ummz),
and Palawan (fmnh).

Habitat — Poorly known; apparently found
only in lowland areas (Sanborn, 1952). Found in
secondary forest, plantations, gardens, scrub, and
grassland on Borneo (Payne et al., 1985).

Status — Abundant in agricultural areas.

Comment — Formerly known as R. jalorensis.

Rhynchomys isarogensis Musser and
Freeman, 1981. J. Mammal., 62:154.

Common Name — Isarog shrew-rat.
Distribution — Known only from Mt. Isarog,
Camarines Sur Province, Luzon (fmnh).

Habitat — Primary montane and mossy forest
from 1125 m to 1750 m (Balete & Heaney, in
press; Heaney et al., in press; Rickart et al., 1991 ).

Status — Geographically restricted (Musser &
Freeman, 1981) and threatened by logging. IUCN:
Vulnerable.

Rhynchomys soricoides Thomas, 1895.
Ann. Mag. Nat. Hist., ser. 6, 16:160.

Common Name — Northern Luzon shrew-rat.

Distribution— Known only from Mt. Data,
Benguet Province, Luzon (fmnh).

Habitat — Known only from mossy forest at
2200 m to 2400 m in the Central Cordillera (Ra-
bor, 1955; Sanborn, 1952; Thomas, 1898).

Status — Uncommon in high-elevation mossy
forest.

Sundamys muelleri (Jentink, 1879).
Notes Ley den Mus., 2:16.

Common Name — Great Sunda rat.

Distribution — Southern Burma to Palawan.
Philippine records are from Balabac (usnm), Bus-
uanga (usnm), Culion (fmnh), and Palawan
(fmnh).

Habitat — Second growth and primary forest,
from sea level to 900 m (Sanborn, 1952).

Status — Common in forest habitats from low-
lands to mossy ridge tops.

Tarsomys apoensis Mearns, 1905. Proc.
U.S. Natl. Mus., 28:453.

Common Name — Dusky moss-mouse, Minda-
nao dusky rat.

Distribution — Known only from the highlands
of Mindanao (Bukidnon [fmnh], Davao del Sur
[fmnh], Misamis Occidental [srj, Misamis Ori-
ental [fmnh], and Zamboanga del Norte [uplb]
provinces).

Habitat — From 1550 m to 2400 m in montane
and mossy forest (Musser & Heaney, 1992).

Status — Moderately common in high-eleva-
tion forest.

Tarsomys echinatus Musser and Heaney,
1992. Bull. Am. Mus. Nat. Hist., 21 1:33.

Common Name — Mindanao spiny rat.
Distribution — Known only from Mindanao

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

43

(Bukidnon [Musser, 1994] and South Cotabato
[dmnh] provinces).

Habitat — Poorly known. Apparently restricted
to lowland forest.

Status — Uncertain. Probably formerly wide-
spread in lowland forest on Mindanao, but may
now be greatly affected by habitat destruction.
IUCN: Vulnerable.

Hystricidae — Porcupines

A single endemic species in this distinctive
family occurs in the Philippines.

Hystrix pumila (Gunther, 1879).
Mag. Nat. Hist., ser. 5, 4:106.

Ann.

Tarsomys sp. A

Common Name — Sibuyan giant moss-mouse,
Sibuyan dusky rat.

Distribution — Sibuyan Island only (fmnh).

Habitat — Primary forest from near sea level to
1325 m (Goodman & Ingle, 1993; fmnh).

Status — Stable but highly restricted geograph-
ically and threatened by logging (Goodman & In-
gle, 1993).

Tarsomys sp. B

Common Name — Buffy-collared moss-mouse.

Distribution — Known only from Mt. Kitang-
lad, Bukidnon Province, Mindanao (fmnh).

Habitat — Known only from 2250 m to 2800
m in primary mossy forest (Heaney et al., unpubl.
data).

Status — Moderately common in high-eleva-
tion forest, but probably restricted geographically.

Comment — Assignment of this and the preced-
ing undescribed species to the genus Tarsomys is
tentative. Both species appear to share characters
with both Tarsomys and Limnomys.

Tryphomys adustus Miller, 1910. Proc.
U.S. Natl. Mus., 38:399.

Common Name — Luzon short-nosed rat.

Distribution — Luzon Faunal Region; known
only from Benguet (fmnh), Laguna (usnm), and
Tarlac (usnm).

Habitat — Mossy forest at about 2500 m in the
Central Cordillera (Miller, 1910; Rabor, 1955) and
in the lower parts of Mt. Makiling (ca. 100-350
m) (Barbehenn et al., 1973).

Status — Uncommon but widespread in central
Luzon (Musser, 1982b). IUCN: Vulnerable.

Comment — The genus Tryphomys was re-
viewed and redefined by Musser and Newcomb
(1983) and Musser and Heaney (1992).

Common Name — Palawan porcupine.

Distribution — Palawan Faunal Region. Re-
corded on Busuanga (fmnh) and Palawan (fmnh).

Habitat — Lowland secondary and primary for-
est (Hoogstraal, 1951).

Status — Locally common to uncommon
(Hoogstraal, 1951).

Carnivora

Felidae— Cats

Only one species of true cat occurs in the Phil-
ippines (Fig. 7); other species sometimes called
cats are actually members of the family Viverri-
dae. The one species is widespread in Southeast
Asia, and has a limited distribution in the Philip-
pines. Feral domestic cats occasionally live in for-
ested areas on Luzon and perhaps elsewhere.

Prionailurus bengalensis (Kerr, 1792). In
Linnaeus, Anim. Kingdom, 1:151.

Common Name — Leopard cat.

Distribution — Siberia to Pakistan and Bali. In
the Philippines, documented only on Negros
(fmnh) and Palawan (fmnh), and reported from
Busuanga, Cebu, and Panay (Taylor, 1934; Timm
& Birney, 1980).

Habitat — Recorded in agricultural habitats and
forest from sea level to about 1500 m (Rabor,
1986).

Status — Uncommon but widespread; heavily
hunted and declining in the Philippines.

Comment — Prionalurus is now recognized as a
genus, rather than a subgenus of Felis (Wozen-
craft, 1993). Populations from the Palawan Faunal
Region and from the Negros-Panay Faunal Re-
gion are being described as distinct subspecies
(Groves, pers. comm.).

44

FIELDIANA: ZOOLOGY

Fig. 7. Phonailurus bengalensis. the leopoard cat; a rarely seen cat that is widespread in Southeast Asia but
restricted in the Philippines to the Palawan and Negros-Panay faunal regions.

Mustelidae — Weasels, Otters, and Badgers

Only two species of this family occur in the
Philippines, one otter and one stink-badger; the
latter is a Philippine endemic.

Amblonyx cinereus (Illiger, 1815). Abh.
Phys. Klasse K. Preuss. Akad. Wiss.,
[1804-1811] p. 99.

Common Name — Oriental small-clawed otter.
Distribution — India to Taiwan and Java; Phil-
ippine records are from Palawan Island only

(I \1NH).

Habitat — Along coastal rivers and bays (San-
born, 1952).

Status — Widespread, but limited distribution
within the Philippines; probably uncommon.

Comment — Amblonyx was recognized as a val-
id genus, rather than a subgenus of Aonyx, by
Wozencraft (1993).

Mydaus marchei (Huet, 1887). Le Natur-
aliste, ser. 2, 9(13): 149-151.

Common Name — Palawan stink-badger.

Distribution — Palawan Faunal Region. Re-
corded only on Busuanga (fmnh), Calauit (Dans,
unpubl. data), and Palawan (usnm).

Habitat — Mixed grassland and second-growth
forest (Hoogstraal, 1951; Rabor, 1986).

Status — Geographically restricted and locally
moderately common to uncommon in secondary
and primary lowland forest. IUCN: Vulnerable.

Herpestidae — Mongooses

A single member of this widespread family oc-
curs in the Philippines.

Herpestes brachvurus Gray, 1837. Proc.
Zool. Soc. Lond. [1836] p. 88.

Common Name — Short-tailed mongoose.
Distribution — Malay Peninsula to Borneo and

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

45

Palawan. Philippine records are from Palawan
(fmnh) and Busuanga islands only.

Habitat — Uusually found near rivers (Rabor,
1986). On Borneo, it occurs in lowland primary
and secondary forest, as well as on plantations
and in gardens (Payne et al., 1985).

Status — Widespread, probably moderately
common, but current status poorly known.

Viverridae — Civets

Three species of this diverse family occur in
the Philippines. Two species are widespread in the
Philippines and are the only mammalian carni-
vores to occur in most of the country. None of
the three is endemic.

Arctictis binturong (Raffles, 1821).
Trans. Linn. Soc. Lond. 13:253.

Common Name — Binturong.

Distribution — Northern Burma and Yunnan to
Sumatra, Java, and Borneo. In the Philippines,
known only from Palawan Island (amnh).

Habitat — Primary and secondary lowland for-
est from sea level to 200 m (Rabor, 1986).

Status — Widespread, but Philippine popula-
tions restricted and uncommon.

Comment — Palawan Island specimens were
originally described as a distinct species. A. whitei
(Allen, 1910), and are periodically recognized as
such.

Paradoxurus hermaphroditus (Pallas,
1777). In Schreber, Die Saugethiere,
3(25):426.

Common Name — Common palm civet.

Distribution — Sri Lanka to Hainan and the
Lesser Sunda Islands; found throughout the Phil-
ippines. Specimens from Balabac (usnm), Bus-
uanga (fmnh), Camiguin (fmnh), Culion (fmnh),
Leyte (usnm), Luzon (Bataan [usnm], Cagayan
[ummz], Camarines Sur [fmnh], Ilocos Norte
[usnm], Isabela [amnh], Laguna [amnh], Pampan-
ga [usnm], Rizal [amnh], and Sorsogon [dmnh]
provinces), Marinduque (pnm), Mindanao (Agu-
san del Norte [su], Bukidnon [fmnh], Davao del
Norte [fmnh], Davao del Sur [fmnh], Lanao del
Sur [dmnh], Maguindanao [fmnh], Misamis Oc-
cidental [usnm], South Cotabato [fmnh], Surigao

del Norte [su], Surigao del Sur [dmnh], Zam-
boanga del Norte [fmnh], and Zamboanga del Sur
[dmnh]), Negros (fmnh), Palawan (fmnh), Sanga-
sanga (dmnh), and Sibuyan (fmnh). Also reported
from Catanduanes (Heaney et al., 1991), Biliran,
Maripipi (Rickart et al., 1993), and Panay (Timm
& Birney, 1980; Lastimosa, pers. coram.).

Habitat — Recorded in agricultural and forest-
ed areas from sea level up to at least 2400 m
(Balete & Heaney, in press; Heaney et al., 1991,
in press; Hoogstraal, 1951; Rabor, 1986; Thomas,
1898).

Status — Common and geographically wide-
spread.

Viverra tangalunga Gray, 1832. Proc.
Zool. Soc. Lond., p. 63.

Common Name — Malay civet, tangalung.

Distribution — Malay Peninsula to Sulawesi
and Amboina; found throughout the Philippines.
Specimens from Bohol (fmnh), Busuanga (fnmh),
Culion (fmnh), Leyte (usnm), Luzon (Abra
[fmnh], Bataan [amnh], Camarines Sur [usnm],
Cagayan [ummz], and Isabela [amnh] provinces),
Mindanao (Davao del Norte [fmnh], Davao del
Sur [fmnh], Lanao del Sur [usnm], Maguindanao
[fmnh], Misamis Occidental [usnm], and Misamis
Oriental [fmnh] provinces), Mindoro (usnm), Ne-
gros (usnm), Palawan (usnm), Samar (fmnh), and
Sibuyan (fmnh). Also reported from Camiguin
(Heaney & Tabaranza, 1995), Catanduanes (Hea-
ney et al., 1991), Panay (Timm & Birney, 1980;
Lastimosa, pers. comm.), and Siguijor (Timm &
Birney, 1980).

Habitat — In primary and secondary lowland,
montane, and mossy forest from sea level to at
least 1200 m (Heaney et al., in press; Rabor, 1955;
Rickart et al., 1993).

Status — Widespread in Asia. Moderately com-
mon in forest, rare elsewhere.

Artiodactyla

Suidae — Pigs

Three speces of wild pigs occur in the Philip-
pines (Groves, 1997), two of which are endemic.

46

FIELDIANA: ZOOLOGY

Sus barbatus Miiller, 1838. Tijdschr. Nat.
Gesch. Physiol., 5:149.

Common Name — Bearded pig.

Distribution — Malay Peninsula to Borneo and
Palawan Faunal Region (Caldecott et al., 1993).
Found on Busuanga (fmnh) and Palawan (fmnh).
Also reported from Balabac (Taylor, 1934) and
Bugsuc, Calauit, Culion, Coron, and adjacent is-
lands (Oliver, unpubl. data).

Habitat — Originally occurred from sea level to
highest peaks, in primary and secondary forest
(Rabor, 1986).

Status — Widespread and locally common, but
heavily hunted and declining (Caldecott et al.,
1993; Oliver, 1992).

Comment — The subspecies S. b. ahoenobarbus
is restricted to the Palawan Faunal Region and is
listed as a threatened taxon by the IUCN.

Sus cebifrons Heude, 1888. Mem. Hist.
Nat. Emp. Chin., 2, pi. 17, Fig. 5.

Common Name — Visayan warty pig.

Distribution — Negros-Panay Faunal Region
only. Specimens have been taken from Masbate
(fmnh) and Negros (fmnh). It is also reported
from Cebu, Guimaras, and Panay Islands (Oliver,
1992; Oliver et al., 1993b).

Habitat — Originally in primary and secondary
forest from sea level to mossy forest at 1600 m;
now found only above 800 m.

Status — Extinct on Cebu and Guimaras.
Heavily hunted and increasingly rare; now hy-
bridizing with domestic pigs (Oliver, 1992).
IUCN: Critically endangered.

Comment — Recognized as a distinct species by
Groves and Grubb (1993) and Grubb (1993).

Sus philippensis Nehring, 1886. Sber.
Ges. Naturf. Fr., Berlin, p. 83.

Common Name — Philippine warty pig.

Distribution — Biliran (ummz), Catanduanes

(fmnh), Leyte (usnm), Luzon (Abra [fmnh], and

Rizal [fmnh] provinces), Mindanao (Bukidnon

fmnh], Davao del Norte [fmnh], Davao del Sur

fmnh], Maguindanao [fmnh]. North Cotabato

fmnh], South Cotabato [fmnh], and Zambonanga

del Norte [fmnh] provinces), and Mindoro (fmnh).

Also reported from Basilan. Samar (Oliver, 1992).

and CamiL'um <Heane\ & Tabaranza. unpubl.
data).

Habitat — Formerly abundant from sea level to
at least 28(K) m. in virtually all habitats (Rabor.
1986); now common only in remote forests (Dan
ielsen et al., 1994; Heane) et al.. 1991. in press)

Status — Heavily hunted, declining rapidly
(Garcia & Deocampo. 1995). Also threatened by
hybridization in some areas Extinct on Mann-
duque (Oliver. 1992).

Comment — Recognized as a distinct species by
Grubb (1993).

Tragulidae — Mouse-deer

A single species of this unusual family barely
enters the Philippines.

Tragulus napu (F Cuvier, 1822). In E.
Geoffroy and F. Cuvier, Hist. Nat. Mam-
miferes, part 2, 4(37): 4 pp.

Common Name — Greater mouse-deer.

Distribution — Southern Indochina to Java and
Borneo. In the Philippines, recorded only on Bal-
abac (fmnh) and adjacent small islands, including
Bugsuc and Ramos (Oliver, unpubl. data).

Habitat — Primary and secondary forest and
scrub (Hoogstraal, 1951; Rabor, 1986).

Status — Species widespread and common.
The Philippine population is highly restricted but
reported to be locally common in some areas in
1993, despite continuous heavy hunting pressure
(Oliver et al., unpubl. data).

Cervidae — Deer

Three species of deer occur in the Philippines,
all of which are endemic to the country. A ten-
tative report of a possibly introduced population
of Cenus nippon on Jolo (Grubb & Groves.
1983) has yet to be confrimed (Heane) et al ..
1987).

Axis calamianensis (Heude, 188S). Mem
Hist. Nat. Emp. Chin. 2:49.

Common Name — Calamian hog -deer.
Distribution — Palawan Faunal Region onlj

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

47

Recorded on Busuanga (fmnh) and Culion
(fmnh).

Habitat — Grasslands and second growth
(Hoogstraal, 1951).

Status — Vulnerable because of a very limited
range and continued hunting pressure (Oliver,
1994). IUCN: Endangered. CITES: Appendix I.
U.S. ESA: Endangered.

Comment — Regarded as a valid species by
Grubb (1993) and Corbet and Hill (1992), with all
authors recognizing Axis as a genus rather than a
subgenus of Cervus.

Cervus alfredi Sclater, 1876. Proc. Zool.
Soc. Lond., p. 381.

Common Name — Visayan spotted deer.

Distribution — Philippines only; Cebu, Gui-
maras, Masbate, Negros, and Panay islands (Oli-
ver, 1994; Oliver et al., 1992).

Habitat — Formerly from sea level to at least
1500 m in primary forest and second growth.

Status — Geographically restricted and rare.
Now extinct on Cebu, Guimaras, and probably
Masbate. Heavily hunted, severely endangered
(Cox, 1987; Evans et al., 1993; Oliver, 1994; Ol-
iver et al., 1992). IUCN: Endangered. U.S. ESA:
Endangered.

Comment — Recognized as a distinct species by
Grubb and Groves (1983).

Cervus mariannus Desmarest, 1822.
Mammalogie. In Encycl. Meth., 2:436.

Common Name — Philippine brown deer.

Distribution — Originally restricted to the Phil-
ippines but introduced into the Marianna Islands.
Occurs throughout most of the Philippines except
the Negros-Panay Faunal Region, the Babuyan/
Batanes groups, the Palawan Fanual Region, and
the Sulu Faunal Region (Oliver et al., 1992). Re-
corded on Basilan (fmnh), Catanduanes (fmnh),
Leyte (usnm), Luzon (Isabela [ummz] Province),
and Mindanao (Bukidnon [fmnh], Davao del Nor-
te [fmnh], Davao del Sur [fmnh], and South Cot-
abato [fmnh] provinces). Extinct on Biliran (Rick-
art et al., 1993). Also reported from Mindanao
(Augusan del Norte Province) by Sanborn (1953).

Habitat — Formerly from sea level to at least
2900 m in primary and secondary forest (Heaney
et al., in press; Rabor, 1986; Sanborn, 1952; Tay-
lor, 1934).

48

Status — Locally common in isolated areas, but
heavily hunted and declining (e.g, Danielsen et
al., 1994). Local extinctions have been reported
on Catanduanes and Biliran (Heaney et al., 1991;
Rickart et al., 1993).

Comment — Substantial variation in size and
color often exist in limited areas. Further taxo-
nomic study is needed.

Bovidae — Cattle

A single species in this family is native to the
Philippines. It is one of the most seriously endan-
gered species in the family.

Bubalus mindorensis (Heude, 1888).
Mem. Hist. Nat. Emp. Chin., 2:4.

Common Name — Tamaraw, Mindoro dwarf
buffalo.

Distribution — Endemic to the Mindoro Faunal
Region; found in remote areas on Mindoro only
(fmnh).

Habitat — Originally from sea level to high
peaks on Mindoro, but now confined to a few re-
mote areas in rough terrain. Probably prefers sec-
ond growth and mixed forest/grassland (Kuehn.
1986; Custodio et al., 1996).

Status — Rare and geographically restricted;
severely endangered and declining (Custodio et
al., 1996; Oliver, 1994). IUCN: Endangered.
CITES: Appendix I. U.S. ESA: Endangered.

Comment — Groves (1969) assigned the tama-
raw to the subgenus Bubalus, rather than to Anoa.

Cetacea

Twenty-one species of cetaceans have been re-
liably documented from Philippine waters (Alava
et al., 1993; Tan, 1995). Additionally, there are
unconfirmed or suspected occurrences of the fol-
lowing species (Leatherwood et al., 1992): Ba-
laenoptera musculus, Delphinus delphis, Hype-
roodon sp., Mesoplodon ginkgodens, and Sousa
chinensis.

Balaenopteridae — Rorquals

Three of the six species in this family occur in
the Philippines.

FIELDIANA: ZOOLOGY

Balaenoptera acutorostrata Lacepede,
1804. Hist. Nat. Cetacees, p. 134.

Common Name — Minke whale.

Distribution — Worldwide. Very rare in some
tropical pelagic areas such as the eastern tropical
Pacific (Jefferson et al., 1994). In the Philippines,
a stranding was reported by Herre (1925) at Ba-
coor, Cavite Province (Luzon), but individuals
have yet to be sighted at sea.

Habitat — Coastal, inshore, and offshore areas.

Status — Apparently uncommon (Klinowska,
1991). CITES: Appendix I. IUCN: Insufficiently
known.

Balaenoptera edeni Anderson, 1878.
Anat. Zool. Res., Yunnan, p. 551, pi. 44.

Common Name — Bryde's whale, tropical
whale.

Distribution — Tropical and subtropical zones.
This species generally does not move beyond 40°
in either hemisphere (Jefferson et al., 1994). In
the Philippines, sighted in Panay Gulf and the
eastern Sulu Sea in May, 1995 (Dolar, unpubl.
data). Skeleton specimens are housed in suml.

Habitat — Coastal and offshore waters.

Status — Scarce and poorly known (Klinows-
ka, 1991). This species is being exploited by hunt-
ers from Pamilacan, Bohol, and Camiguin (north
of Mindanao). Hunting grounds are the Bohol Sea
(sometimes called the Mindanao Sea) and the
eastern Sulu Sea (Dolar et al., 1994; Leatherwood
et al., 1992). CITES: Appendix I. IUCN: Insuffi-
ciently known.

Comment — Philippine Bryde's whales may be
the small coastal form of Bryde's whales found in
other regions. Morphometric studies showed that
they are relatively smaller than those found in oth-
er regions (Perrin et al.. 1996b). Genetic studies
are currently being done to investigate the validity
of this group as a separate species. Preliminary
results indicate that the Southeast Asian group is
genetically different from all other populations of
Bryde's whale and may deserve recognition as a
separate species (Dizon et al., 1996).

Megaptera novaeangliae (Borowski,
1781). Gemein. Naturgesch. Thier., 2(1):
21.

Common Name — Humpback whale.
Distribution — Worldwide. Sighted near south-

eastern Palawan, northwest of Luzon and off
western Mindanao (Slijper et al., 1964).

Habitat — Feed and breed in coastal waters and
migrate from the tropics to polar and subpolar
regions, reaching the ice edges in both hemi-
spheres (Jefferson et al., 1994).

Status — Generally rare (Klinowska, 1991).
This species has not been sighted in Philippine
waters in recent years. CITES: Appendix I. IUCN:
Vulnerable.

Delphinidae — Ocean Dolphins

Twelve of the 32 species in this family occur
in the Philippines. None is endemic.

Feresa attenuata Gray, 1875. J. Mus.
Godeffroy (Hamburg), 8:184.

Common Name — Pygmy killer whale.

Distribution — Circumtropical and subtropical
waters, generally not ranging beyond 40°N and
35°S (Jefferson et al., 1994). This species has
been sighted in the Bohol Sea, Bohol Strait, Tanon
Strait, and eastern Sulu Sea (Dolar & Perrin,
1996; Dolar & Wood, 1993; Leatherwood et al.,
1992). Specimens are from the eastern Sulu Sea
(suml).

Habitat — Oceanic.

Status — Moderately common in Philippine
waters. CITES: Appendix II. IUCN: Insufficiently
known (Klinowska, 1991).

Globicephala macrorhynchus Gray,
1846. Zool. Voy. H.M.S.' "Erebus" and
"Terror", 1:33.

Common Name — Short-finned pilot whale.

Distribution — Warm temperate to tropical wa-
ters, usually between 50°N and 40°S. Sighted in
the Sulu and Bohol seas and Tanon Strait (Dolar
& Wood, 1993; Leatherwood et al., 1992). Stand-
ings recorded in Calategas, Narra, Palawan, and
Binmaley, Pangasinan Province, Luzon (Leather-
wood et al., 1992). Specimens are in the suml.
uplb, and AMNH.

Habitat — Usually deep offshore areas. In the
Philippines, often found over or near steep slopes
(Dolar & Perrin, 1996).

Status — Common in the Philippines. CITES:

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

49

Appendix II. IUCN: Insufficiently known (Kli-
nowska, 1991).

Comment — In the eastern Pacific, often found
associated with other species (e.g., bottlenose, Pa-
cific white-sided, and Risso's dolphins and sperm
whales; Jefferson et al., 1994). In the Sulu Sea,
Philippines, found with Fraser's dolphins in 85%
of the total sightings (Dolar & Perrin, 1996).

Grampus griseus (G. Cuvier, 1812). Ann.
Mus. Hist. Nat. Paris, 19:13.

Common Name — Risso's dolphin.

Distribution — Worldwide. Sighted in the Tan-
on Strait, Visayan Sea, Bohol Sea, Cebu Strait,
Camotes Sea, eastern Sulu Sea, Celebes Sea, east-
ern China Sea (Dolar & Perrin, 1996; Dolar &
Wood, 1993; Hammond & Leatherwood, 1984;
Leatherwood et al., 1992), and southern Sulu Sea
(Dolar & Perrin, unpubl. data). Specimen known
from eastern Sulu Sea (suml).

Habitat — Coastal and offshore waters.

Status — Moderately common in the Philip-
pines. CITES: Appendix II. IUCN: Insufficiently
known (Klinowska, 1991).

Lagenodelphis hosei Fraser, 1956. Sara-
wak Mus. J., n.s., 8(7):496.

Common Name — Fraser's dolphin.

Distribution — Pantropical, largely between
30°N and 30°S. Distributed widely in the Philip-
pines. Sighted in the Bohol Sea, Cebu Strait, Sulu
Sea, and Celebes Sea (Dolar & Perrin, 1996; Do-
lar & Wood, 1993; Leatherwood et al., 1992;
Hammon & Leatherwood, 1984). Specimens are
from the eastern Sulu Sea (suml).

Habitat — Oceanic but can be seen nearshore
in areas where deep water approaches the coast
as in the Philippines (Dolar & Perrin 1996; Jef-
ferson et al., 1994).

Status — Very common in the Philippines.
CITES: Appendix II. IUCN: Insufficiently known
(Klinowska, 1991).

Orcinus orca (Linnaeus, 1758). Syst.
Nat., 10th ed., 1:77.

Common Name — Killer whale, orca.
Distribution — Worldwide. Sighted only once,
in the Sulu Sea (Dolar & Perrin, 1996).

Habitat — Found in almost any marine region,
including nearshore, offshore, and oceanic waters.
The species is known to ascend rivers (Jefferson
et al., 1994).

Status — Apparently rare in the Philippines.
CITES: Appendix II. IUCN: Insufficiently known
(Klinowska, 1991).

Peponocephala electra (Gray, 1846).
Zool. Voy. H.M.S. "Erebus" and "Ter-
ror", 1:35.

Common Name — Melon-headed whale.

Distribution — Tropical and subtropical ocean-
ic waters between 40°N and 35°S (Jefferson et al.,
1994). In the Philippines this species is seen in
relatively shallow waters. It is locally common in
the Tanon Strait and near Siquijor Island. Sighted
in the eastern Sulu and Bohol seas (Dolar & Per-
rin, 1996; Dolar & Wood, 1993; Leatherwood et
al., 1992) and Tanon Strait (Dolar & Perrin, un-
publ. data) and Cebu Strait (Hammond & Leath-
erwood, 1984).

Habitat — Coastal and oceanic waters. In the
Philippines only seen in relatively shallow waters
(^ 200 fathoms) close to shore.

Status — Locally common in the Philippines.
CITES: Appendix II. IUCN: Insufficiently known
(Klinowska, 1991).

Pseudorca crassidens (Owen, 1846).
Hist. Brit. Foss. Mamm. Birds, p. 516,
Fig. 213.

Common Name — False killer whale.

Distribution — Tropical to warm temperate
zones. Do not range beyond 50° in either hemi-
sphere. Sighted in the Philippine Sea (northeast
and east of Luzon) and the South China Sea
(northwest of Luzon; Ferrin et al., 1996a).

Habitat — Deep offshore waters. In the Philip-
pines, the most inshore locality was approximate-
ly 80 km northeast of San Miguel Bay, Luzon
(Perrin et al., 1996a).

Status — Insufficiently known, probably un-
common in the Philippines. CITES: Appendix II.
IUCN: Insufficiently known (Klinowska, 1991).

Stenela attenuata (Gray, 1846). Zool.
Voy. H.M.S. "Erebus" and "Terror", 1:
44.

Common Name — Pantropical spotted dophin.
Distribution — Tropical and subtropical zones.

50

FIELDIANA: ZOOLOGY

Found in all oceans bet wen 40°N and 40°S, al-
though more abundant in lower latitudes (Jeffer-
son et al., 1994). Sighted in the Sulu Sea, South
China Sea (west of Palawan), Tanon Strait, Cebu
Strait, Bohol Sea, Batangas Bay, Verde Island
Passage, and Celebes Sea (Dolar et al., 1994; Do-
lar & Perrin, 1996; Dolar & Wood, 1993; Ham-
mond & Leatherwood, 1984; Leatherwood et al.,
1992). Specimens are from the eastern Sulu Sea
and Bohol Sea (suml).

Habitat — Nearshore and offshore waters.

Status — Abundant in the Philippines. CITES:
Appendix II. 1UCN: Insufficiently known (Kli-
nowska, 1991).

Stenella coeruleoalba (Meyen, 1833).
Nova Acta Acad. Caes. Nat. Curios.,
16(2):609, pi. 43.

Common Name — Striped dolphin.

Distribution — Primarily in warm waters, but
also sometimes seen in temperate regions (be-
tween 50°N and 40°S) (Jefferson et al., 1994).
Sighted in the Philippine Sea (northeast of Luzon)
and the South China Sea (northwest of Luzon and
west of Batanes; Perrin et al., 1996a).

Habitat — Oceanic waters. Seen near shore
only when deep water approaches the coast (Jef-
ferson et al., 1994).

Status — Insufficiently known, probably un-
common in the Philippines. CITES: Appendix II.
IUCN: Insufficiently known (Klinowska, 1991).

Stenella longirostris (Gray, 1828). Spicil.
Zool., 1:1.

Common Name — Spinner dolphin.

Distribution — Tropical and subtropical zones,
between 40°N and 40°S. Sighted in the China Sea
(west of Palawan), Sulu Sea, Bohol Sea, Tanon
Strait, Cebu Strait, Verde Island Passage, Camotes
Sea, Batangas Bay, and Celebes Sea (Dolar &
Perrin, 1996; Dolar & Wood, 1993; Hammond &
Leatherwood, 1984; Leatherwood et al., 1992).
Specimens are from the Sulu and Bohol seas
(suml).

Habitat — Nearshore and oceanic waters.

Status — Most abundant dolphin in the Philip-
pines. CITES: Appendix II. IUCN: Insufficiently
known (Klinowska, 1991).

Steno bredanensis (Lesson, 1828). Hist.
Nat. Gen. Part. Mamm. Oiseaux, 1:206.

Common Name — Rough-toothed dolphin.

Distribution — Tropical to subtropical, between
40°N and 35°S. Sighted in the Sulu Sea (Dolar,
unpubl. data), accidentally caught in a fishery in
the Celebes Sea (Dolar & Wood, 1993).

Habitat — Deep oceanic waters.

Status — Uncommon in the Philippines.
CITES: Appendix II. IUCN: Insufficiently known
(Klinowska, 1991).

Tursiops truncatus (Montagu, 1821).
Mem. Wernerian Nat. Hist. Soc, 3:75,
pi. 3.

Common Name — Bottlenose dolphin.

Distribution — Tropical and temperate, be-
tween 45°N and 45°S. Sighted in the Sulu Sea,
Bohol Sea, Tanon Strait, Batangas Bay, Verde Is-
land Passage, and Ulugan Bay (Dolar & Perrin,
1996; Dolar & Wood, 1993; Leatherwood et al.,
1992). Locally common in the Panay Gulf and
southern Sulu Sea (Dolar & Perrin, unpubl. data).

Habitat — Often found in nearshore and in-
shore waters. Also found to inhabit pelagic waters
(Jefferson et al., 1994).

Status — Locally common in the Philippines.
CITES: Appendix II. IUCN: Insufficiently known
(Klinowska, 1991).

Kogiidae — Dwarf and Pygmy Sperm Whales

There are two species in this family and both
occur in the Philippines.

Kogia breviceps (de Blainville, 1838).
Ann. Franc. Etr. Anat. Phys., 2:337.

Common Name — Pygmy sperm whale.

Distribution — Tropical to warm temperate
zones of all oceans. This species has been sighted
in Tanon Strait, and a skeleton was obtained from
Pamilacan, Bohol (Dolar & Perrin, 1996; Dolar &
Wood, 1993; Leatherwood et al., 1992).

Habitat — Often found over and near the con-
tinental slope.

Status — Uncommon in the Philippines.
CITES: Appendix II. IUCN: Insufficiently known
(Klinowska, 1991).

iHEANEY ET AL.: MAMMALS OF THE PHILIPPINES

51

Kogia simus (Owen, 1866). Trans. Zool.
Soc. Lond., 6(1):30, pis. 10-14.

Common Name — Dwarf sperm whale.

Distribution — Appears to be distributed wide-
ly in tropical to warm temperate areas. Sighted in
the Tanon Strait, Bohol Sea, and eastern Sulu Sea
(Dolar & Perrin, 1996; Dolar & Wood, 1993;
Leatherwood et al., 1992). Specimens are from
the Sulu Sea and Bohol Sea (suml).

Habitat — Appear to be distributed largely off-
shore. Commonly found in inshore waters in Tan-
on Strait (Dolar & Perrin, unpubl. data).

Status — Locally common in the Philippines.
CITES: Appendix II. IUCN: Insufficiently known
(Klinowska, 1991).

Comment — Has been observed frequently in
Tanon Strait. Average sighting is between 6-8/day
(10 hr survey time; Dolar & Perrin, 1996).

Physeteridae — Sperm Whales

The sole species in this family is present in the
Philippines.

Physeter catodon Linnaeus, 1758. Syst.
Nat., 10th ed., 1:76.

Common Name — Sperm whale.

Distribution — Worldwide. Sighted in the Sulu,
Bohol, and Celebes seas (Dolar & Perrin, 1996;
Dolar & Wood, 1993; Leatherwood et al, 1992).
A stranding was recorded in Palawan (Leather-
wood et al., 1992). A specimen was taken from
the Sulu Sea (suml).

Habitat — Oceanic, but also found nearshore
where deep water occurs near the coast (Jefferson
et al., 1994).

Status — Commonly seen in some areas.
CITES: Appendix I. U.S. ESA: Endangered.
IUCN: Insufficiently known (Klinowska, 1991).

Ziphiidae — Beaked Whales

Two of the 19 species in this family are known
to occur in the Philippines. Some unidentified
beaked whales have been sighted in the Sulu Sea
(Dolar & Perrin, unpubl. data). It is probable that
these represent one or more additional species.

Mesoplodon densirostris (de Blainville,
1817). Nouv. Diet. Hist. Nat., Nouv. ed.,
9:178.

Common Name — Blainville's beaked whale,
dense beaked whale.

Distribution — Temperate and tropical waters
in all oceans. Sighted in the Bohol and Sulu seas
(Dolar & Perrin, 1996). Specimen taken from the
Bohol Sea (suml).

Habitat — Usually in offshore and deep waters.
Sometimes found in deep inshore waters in the
Philippines.

Status — Moderately common. CITES: Appen-
dix II. IUCN: Insufficiently known.

Ziphius cavirostris G. Cuvier, 1823.
Rech. Oss. Foss., Nouv. ed., 5(1):350.

Common Name — Cuvier 's beaked whale.

Distribution — Widely distributed in offshore
waters of all oceans. In the Philippines, it was
sighted in the south Sulu Sea, east of Mapun (Ca-
gayan de Tawi-tawi) on 8 May 1996 (Dolar &
Perrin, unpubl. data), and one was caught in a
drift gillnet in the eastern Sulu Sea on 10 June
1996 (suml).

Habitat — Deep offshore waters.

Status — Poorly known, probably uncommon.
IUCN: Insufficiently known (Klinowska, 1991).

Phocoenidae — Porpoises

One of the six species in this family occurs in
the Philippines.

Neophocaena phocaenoides (G. Cuvier,
1829). Regne Anim., Nouv. ed., 1:291.

Common Name — Finless porpoise.

Distribution — Indo-pacific waters, tropical to
temperate zones. Sighted in the South China Sea,
northwest of Palawan (Tan, 1995).

Habitat — Generally found in shallow waters,
both in marine and estuarine areas, or even great
distances up some rivers (e.g., the Yangtze River
in China).

Status — Uncommon in the Philippines.
CITES: Appendix II. IUCN: Insufficiently known
(Klinowska, 1991).

52

FIELDIANA: ZOOLOGY

Sirenia

Dugongidae — Dugongs and Sea Cows

There is only one living species in this family.
It formerly occurred throughout much of the Phil-
ippines.

Dugong dugon (Miiller, 1776). Linne's
Vollstand. Natursyst. Suppl., p. 21.

Common Name — Dugong.

Distribution — Tropical coasts of Indian and
Pacific oceans. Existing reports and interviews
suggest that dugongs previously were present
throughout the Philippine Archipelago. Areas in
the Philippines known to have had dugongs in-
clude Mindoro, Zambales, Palawan, Camarines
Norte, Manila, Cebu, Zamboanga, Tacloban, Pa-
nay Island, Sulu Archipelago, Catanduanes, Mas-
bate, Polillo Island, Bicol Region, Iloilo, Balut Is-
land in Sarangani Bay, Palanan in Isabela, Agusan
del Norte, Samar, Aparri, Mati in Davao del Sur,
Misamis, Cuyo, Pangasinan, Bataan, and Cavite
(Yaptinchay, 1994). More recently, this species
has been recorded from Culion (fmnh), El Nido,
(Kataoka, 1987, in Yaptinchay, 1994), Shark Fin
Bay, Taytay, Palawan; Calauit Island and Gutob
Bay, Busuanga (Trono et al., 1993, in Yaptinchay,
1994). Reported in ca. 1991 from northeastern
Luzon (Danielsen et al., 1994). In Romblon Prov-
ince and Bicol (southern Luzon), interview re-
spondents confirmed presence and exploitation of
dugongs (Yaptinchay, 1994). A calf was stranded
in Romblon Province in May 1993 (Yaptinchay,
1994).

Habitat — Shallow tropical seas with abundant
sea-grass.

Status — Has been heavily exploited in the
Philippines, almost to extinction. CITES: Appen-
dix I. IUCN: Vulnerable. U.S. ESA: Endangered.

Comment — Until the late 1970s, dugongs were
reported present in most of the areas mentioned
above. Today, Palawan is the only place in the
Philippines where reports are regularly received
and confirmed (Yaptinchay, 1994).

Discussion and Conclusion

The mammalian fauna of the Philippines is a
remarkable assemblage of species that occur from

the depths of the sea to the tops of cloud-en-
shrouded mountains. The data provided here are
brief and limited, but they provide a clear over-
view of this fauna. Although we refer the reader
to recent summaries of zoogeography and conser-
vation status for details (Heaney, 1986, 1991b,
1993; Heaney et al., 1997), several key features
will be apparent from perusal of this listing.

First, the terrestrial fauna of the Philippines is
divided into a series of centers of endemism.
These centers correspond to the islands that ex-
isted in the Philippines during periods of glacial
development in high-latitude regions and conse-
quent low sea level worldwide. Every one of these
ice-age islands that has been investigated, includ-
ing the tiny islands of Camiguin and Sibuyan,
supports at least two endemic species, and most
islands have many more endemic species (Hea-
ney, 1985b, 1986, 1991b, 1993). The sole excep-
tion to this pattern is Siquijor, which has no en-
demic species, but also has virtually no native
vegetation remaining (Lepiten, 1995).

Second, native terrestrial mammalian species
occur at every elevation and in every type of for-
est habitat that remains in existence in the country
(e.g., Heaney et al., 1989; Rickart, 1993; Rickart
et al., 1991). On the other hand, few native spe-
cies are known to survive in deforested habitats,
and many native species cannot survive in dis-
turbed forest (Heaney, 1993; Heaney & Utzurrum,
1991).

Third, at least 50 species of terrestrial mammals
currently are known to be threatened to varying
degrees, some critically so. Many others that are
poorly known at present are certain to be added
to this number as more information becomes
available (Heaney, 1993; Heaney & Utzurrum,
1991; Oliver & Heaney, 1996; Utzurrum, 1992).
Endangered species occur in every faunal region
and in all types of natural (undisturbed) habitat.
Populations of most endemic species have de-
clined significantly, the vast majority as a result
of forest habitat destruction. It is clear that forest
destruction has been and remains at present the
primary conservation problem on land in the Phil-
ippines. Additionally, overhunting has had a se-
vere impact on most large terrestrial species and
several marine species, and disturbance and de-
struction of caves has had a major impact on
many species of bats (Utzurrum, 1992). Whale
hunting in the Bohol Sea by fishermen from Pam-
ilacan, Bohol, and Camiguin has almost complete-
ly wiped out the Philippine population of Bryde's
whale. This population is part of the Southeast

HEANEY ET AL.: MAMMALS OF THE PHILIPPINES

53

Asian group found to be morphologically and ge-
netically different from all other Bryde's whales
and may deserve recognition as a separate species
(Perrin et al., 1996b; Dizon et al., 1996).

Fourth, the rich marine mammal fauna is as-
sociated with the archipelagic nature of the coun-
try, which has resulted in the usual bathymetric
topography of its marine environment. Small is-
lands are surrounded by very deep oceanic waters
(the Philippines have some of the deepest waters
in the world), and conversely, deep oceanic waters
are often isolated from each other by narrow, shal-
low sills as well as by islands. This situation has
resulted in complex marine habitats and affected
the distribution of cetaceans. It is common to find
cetaceans that are considered to be "offshore oce-
anic species" (e.g., Fraser's dolphin, Lagenodel-
phis hosei) only 30 m off an island in the Phil-
ippines, which makes them "coastal" species as
well. This has also changed the whole suite of
species associations and assemblages usually seen
elsewhere, making the Philippine situation quite
exceptional.

Fifth, the mammalian fauna of many parts of
the Philippines, both marine and terrestrial, re-
mains poorly known. Detailed, extensive, and
quantitative field studies are crucial in enhancing
knowledge of the fauna. Superficial study will not
yield information on rare or elusive species, nor
will it provide the basic data required for long-
term management decisions.

The data presented in this paper paint a brief
picture of one of the most remarkable mammalian
faunas on Earth: giant bushy-tailed rats, dwarf
buffalos, bats with nearly 6-ft. wingspans, bur-
rowing rodents with white stripes, and "flying le-
murs" that neither fly nor are lemurs. These spe-
cies form a remarkable community of animals that
may be viewed as forming a vast natural experi-
ment in the evolution and maintenance of mam-
malian diversity. Uncovering the story of the or-
igin and ecology of this diversity has provided
one of the major incentives to us in carrying out
the research on which much of this synopsis is
based. A part of the picture, however, is recent
and continuing destruction of forests and reefs,
leaving too many species with only a few places
that can sustain them. It is neither an exaggeration
nor unwarranted melodrama to say that unless ef-
fective action is taken soon to protect the native
habitats of these animals, one of the most re-
markable stories of mammalian evolution on
Earth will end, and an irreplaceable part of the
Filipino heritage will be lost forever. It is our hope

that this paper will assist in the process of devel-
oping effective protection for the environment of
the people and the wildlife of the Philippines, sol-
idly based on knowledge, and dedicated to pro-
viding a stable environment for all.

Acknowledgments

This paper represents the cumulative efforts of
hundreds of people over the course of the last 15
years, from all parts of Philippine society: to all
we express our gratitude. For their assistance with
many and various aspects of the terrestrial re-
search, we especially thank the following people:
Ely Alcala, Nonito Antoque, Willy Arce, Glenn
Bueser, Ron Crombie, Binky Dalupan, Finn Dan-
ielsen, Mariano Roy Duya, Dolly Felicitas, Boy-
ing Fernandez, Linda Flores, Tom Gnoske, Steven
Goodman, Linda Gordon, Paul Heideman, Karl
Hutterer, Arne Jensen, Hans Klompen, Maylene
Laranjo, Myrissa Lepiten, Cres Lumhod, Toto
Manamtam, Pacencia Milan, Marisol dG. Pedre-
gosa, Town Peterson, Leonila Raros, Jim Rieger,
Doug Sampson, David Schmidt, Leoning Tagat,
Joe Walsh, David Willard, and the late Luz Au-
zejo and D. S. Rabor. For their help with field
work on marine mammals, we thank the late Steve
Leatherwood, Colin Wood, William Perrin, Joe
Pres Gaudiano, Moonyeen Alava, Francisco Da-
gat, Esther Himoya, Teddy Redoble, Erwin Rom-
mel Dulombal, Rowena Merto, Alfonso Pading,
Ray Visitacion, Gusieppe Chiu, Gary Aries, Flo-
rian Bernado, Lemnuel Aragones, Cindy Hill,
Earth Island tuna boat observers, Jun Paera, Ella
Delfin, Romella Despu, Michel Gutierrez, Louie
Paera, and Ryan and Chin Lai. We also thank the
officials of the Department of Environment and
Natural Resources who have supported and as-
sisted our efforts, especially Alma Ballesfin, Cor-
azon Catibog-Sinha, Mary Jean Caleda, Carlo C.
Custodio, Wilbur Dee, Josie DeLeon, Luz Gon-
zales, Marlynn Mendoza, Samuel Penafiel, and
Wilfrido Pollisco.

Numerous people at universities and a variety
of nongovernmental organizations have given us
aid in our work; we wish to offer special thanks
to all of our colleagues at the Philippine National
Museum, Silliman University, the University of
the Philippines in Los Banos and Diliman, Min-
danao State University and the Iligan Institute of
Technology, the Haribon Foundation, the Visayas
State College of Agriculture, the Smithsonian In-

54

FIELDIANA: ZOOLOGY

stitution, and the American Museum of Natural
History for their good will and cooperation. We
thank Guy Musser and Karl Koopman for their
encouragement and tutoring in the fine arts of
identification and taxonomy, and M. Carleton, P.
Myers, and R. W. Thorington, Jr. for access to
collections and continuing encouragement. Jodi
Sedlock, Guy Musser, Karl Koopman, and Luis
Ruedas gave helpful comments on an earlier draft
of the manuscript. We thank Ron Crombie, Laurie
Wilkins, and C. A. Ross for access to specimens
and unpublished data. We thank Jodi Sedlock,
Emily McGowan, and Clara Simpson for prepar-
ing the maps and figures, M. Morales, J. Phelps,
M. Solomon, and especially W. T. Stanley for as-
sistance at the Field Museum, and M. Pannell for
her attention to a thousand details in having this
paper become reality. Funding for marine mam-
mal surveys was provided by the World Wildlife
Fund through the Haribon Foundation, the Foun-
dation for the Philippine Environment, the U.S.
National Science Foundation (International Co-
operative Project), the Earth Trust (U.S.), Earth
Island (U.S.), the International Foundation for
Science (Sweden), Greenpeace (U.S.), the Con-
vention for Migratory Species (Germany), Ocean
Park (Hong Kong), and the Whale and Dolphin
Society (U.K.). Terrestrial studies have been sup-
ported by the U.S. National Science Foundation
(BSR-85 14223), the John D. and Catherine T
MacArthur Foundation (90-09272A), and the
Marshall Field, Barbara Brown, and Ellen Thorne
Smith funds of the Field Museum.

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