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Zoology

NEW SERIES, NO. 106

The Mammals and Birds of Camiguin Island,
Philippines, a Distinctive Center of Biodiversity

Lawrence R. Heaney, Editor

April 5, 2006
Publication 1537

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

UNIVERSITY I
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Frontispiece

Two species of mammals and one species of bird live only on Camiguin Island: Apomys sp. (described
herein; center left), Bullimus gamay (described by Rickart et al. in 2002; lower right), and Loriculus sp.
(described herein; upper center). The distinctive volcanic peaks of Camiguin form the background.

8'OLOGY UBRAfly

OCT 2 7 2006

FIELDIANA

Zoology

NEW SERIES, NO. 106

The Mammals and Birds of Camiguin Island,
Philippines, a Distinctive Center of Biodiversity

Lawrence R. Heaney, Editor

Division of Mammals, Department of Zoology
Field Museum of Natural History
1400 South Lake Shore Drive
Chicago, Illinois 60605-2496

Accepted 16 April 2004
Published April 5, 2006
Publication 1537

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 2006 Field Museum of Natural History

ISSN 0015-0754

PRINTED IN THE UNITED STATES OF AMERICA

Contents

1. Mammal and Land Bird Studies on Camiguin Island, Philippines: Background and

Conservation Priorities 1

Laurence R. Heaney and Bias R. Tabaranza, Jr.

2. A New Species of Forest Mouse, Genus Apomys (Mammalia: Rodentia: Muridae), from

Camiguin Island, Philippines 14

Lawrence R. Heaney and Bias R. Tabaranza, Jr.

3. Synopsis and Biogeography of the Mammals of Camiguin Island, Philippines 28

Lawrence R. Heaney, Bias R. Tabaranza, Jr., Natalie Rigertas, and Danilo S. Balete

4. A New Species of Hanging-Parrot (Aves: Psittacidae: Loriculus) from Camiguin Island,

Philippines 49

Jose G. Tello, Jacob F. Degner, John M. Bates, and David E. Willard

5. An Annotated Checklist of the Land Birds of Camiguin Island, Philippines 58

Danilo S. Balete, Bias R. Tabaranza, Jr., and Lawrence R. Heaney

List of Figures (abbreviated titles)

Frontispiece. Color drawing of three endemic species from Camiguin Island (one bird,

two rodents) i

Chapter 1.

1. Photo of Camiguin Island 2

2. Map of Camiguin Island, showing extent of forest and park boundaries 3

3. Map of Camiguin island, showing elevational isoclines and site locations 4

Chapter 2.

1 . Phylogeny of Apomys 17

2. Photo of adult Apomys camiguinensis 18

3. Ventral surface of hind feet of Apomys 20

4. Photos of crania of four species 21

5. SEMs of anterior portions of skulls 22

6. SEMs of posterior portions of skulls 23

7. SEMs of maxillary toothrows 24

8. SEMs of mandibular toothrows 24

Chapter 3.

1. Elevational ranges of non-volant small mammals 31

2. Elevational ranges of fruit bats 34

3. Species accumulation curve 44

Chapter 4.

1 . Map of the central and southern Philippines 50

2. Results of the Principal Components Analysis of morphological data 55

Chapter 5.
No figures

List of Tables (abbreviated titles)

Chapter 1.
None

Chapter 2.

1. Cranial and external measurements of Apomys 16

Chapter 3.

1. Numbers of nonvolant small mammals, by site 32

2. External and cranial measurements of shrews 33

3. Numbers of fruit bats, by site 34

4. External and cranial measurements of fruit bats 35

5. External and cranial measurements of micro bats 38

6. External and cranial measurements of murid rodents 41

Chapter 4.

1. Results of analysis of variance, by sex 53

2. Results of analysis of variance, by taxon, for males 54

3. Results of the principal components analysis 54

Chapter 5.
None

Preface

The Philippine Islands have been recognized
for over a century as having a fauna that is
characterized by large numbers of endemic and
highly distinctive species. For most of this
period, the mammals and birds were thought to
be well known, based on studies conducted in the
period 1880-1950. However, the description of
new species during the 1980s implied that
diversity was higher than had been believed
and suggested that there might be additional
localized centers of endemism yet to be found. In
addition, it was during the 1980s that the extent
of rain-forest habitat destruction became appar-
ent and increased the need for extensive and
detailed documentation of biological diversity to
guide development and conservation for the
benefit of both wildlife and the human popula-
tion of the Philippines.

With this in mind, the authors of this volume
and their colleagues began investigations of
places likely to be of special interest and initiated
a series of publications in Fieldiana: Zoology to
document the findings. The field studies have
been conducted in a standardized fashion to
allow direct and meaningful comparison between
study areas so that geographic patterns of species
richness between islands and mountain ranges
and along elevational gradients could be docu-
mented. However, older museum collections,
which often were unreported in the published
literature, have also been utilized, increasing the
rate of our progress in understanding the re-
markable mammal and bird faunas of the
Philippines. The first publications in Fieldiana
concerned the mammals of Leyte and nearby
islands north of Mindanao, the birds and
mammals of Mt. Isarog National Park in
southern Luzon, and the birds of Sibuyan Island.
Plans are in place for similar publications on the
mammals and birds of the Kitanglad Range of
northern Mindanao and the mammals of Sibu-
yan Island, and others are likely to follow.

This volume contains the results of our studies
of Camiguin Island, a small island that lies just
north of central Mindanao. It is easily over-
looked on a map of the Philippines, yet, as
shown here, the island supports an endemic
species of small parrot and two endemic species
of small mammals, all discovered in the course of
our studies. Announcements in the Philippine
press of the discovery of the mammals in 1994
and 1995 played an important role in encourag-
ing the declaration of the remaining rain forest
on Camiguin as a national park, a movement
that is continuing to gain support. It is our hope
that this publication will help guide the planning
for this protected area and its management when
it has been formally initiated. The data contained
here constitute a baseline from which changes
may be measured in the future and also make
clear how much additional study is needed.
Although Camiguin is a small island, it repre-
sents an ideal natural laboratory in which to
investigate many aspects of the evolution and
conservation of biological diversity.

The assistance of many individuals is acknowl-
edged in each of the chapters that follow, but we
must give special recognition here to the
Philippine Department of Environment and
Natural Resources for providing both permits
and encouragement; we especially thank Dr.
Angel C. Alcala, Dr. Corazon Catibog-Sinha,
Mr. Carlo C. Custodio, Atty. Wilfredo Pollisco,
and Dr. Mundita Lim for their steadfast support.
This project has benefited greatly from the
financial support provided by the John D. and
Catherine T. MacArthur Foundation, from the
Ellen Thorne Smith and Marshall Field Funds of
the Field Museum, and especially the Barbara
Brown Fund for Mammal Research of the Field
Museum.

L. R. Heaney

November 2005
Chicago, Illinois

FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, P. vii

Mammal and Land Bird Studies on Camiguin Island,
Philippines: Background and Conservation Priorities

Lawrence R. Heaney and Bias R. Tabaranza, Jr.

Abstract

Camiguin Island, with an area of ca. 265 km' and maximum elevation of ca. 1620 m, lies
about 10 km north of Mindanao but is isolated from Mindanao by a deep (385 m) channel. It
originated from volcanic activity as a dryland island not earlier than 1 million years ago, but
most growth of the island has occurred within the past 340,000 years. Current landforms are
dominated by large, scenic volcanic peaks, several of which are active. Lowland rain forest
originally occurred up to about 1100 m elevation, with montane rain forest from 1100 m to
about 1350 m and mossy forest from 1350 m to the peaks. By the mid-1990s, deforestation
had removed most natural vegetation below about 600 m, with degree of disturbance to forest
decreasing with elevation and ending at about 1250 m. The climate is tropical, with rainfall of
2-3 m per year in the lowlands and probably about 7.5 m near the peaks. Mammal and/or
bird specimens are available from 18 sites from the 1960s and 1990s; these sites are here
located and described to the extent possible. Given the presence of two endemic species of
mammals (one described in this volume), one endemic bird (described in this volume), and
previously described endemic plants and a frog, Camiguin is one of the smallest but most
distinctive centers of biodiversity in the Philippines and should be a priority site for
conservation. The remaining forest on Camiguin is essential habitat for these unique species,
but it is also essential for watershed protection and control of floods and landslides, and it
contributes significantly to the tourism trade that provides substantial income on the island.
Deforestation for logging and agriculture and overhunting are current threats. A protected
area on the island should include the full range of original habitat diversity, which would
encompass both the existing high-quality forest at upper elevations and also significant tracts
of disturbed but natural lowland forest, especially along rivers and streams, that should be
allowed to regenerate in the future.

Introduction

The mammal and bird faunas of the Philippine
Islands are remarkable for the high total di-
versity of the fauna and especially for the

1 Field Museum of Natural History, 1400 South
Lake Shore Drive, Chicago, IL 60605-2496, U.S.A.

2 Department of Biology, Iligan Institute of Tech-
nology, Mindanao State University, Iligan City,
Lanao del Norte, Philippines.

remarkably large number of endemic species
(e.g., Dickinson et al., 1991; Mittermeier et al.,
1997, 1999; Heaney & Regalado, 1998; Heaney et
al., 1998; Stattersfield et al., 1998). For example,
at least 512 of the 898 species of breeding
terrestrial vertebrates (57%) are endemic to the
Philippines, an unusually high value (Heaney &
Regalado, 1998). Most of the species endemic to
the country occur on the large islands of Luzon,
Mindanao, Mindoro, Negros, and Palawan (e.g.,
Hauge et al., 1986; Heaney, 1986; Heaney et al.,
1998; Collar et al., 1999; Peterson et al., 2000),

FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, PP. 1-13

Fig. 1. Photograph of Camiguin Island from south-southeast, taken on 27 March 1995. All the volcanic peaks
visible on the island are less than 350.000 years old: Mt. Ginsiliban, in the left foreground, is the oldest (see text).

but significant numbers of endemic mammals are
restricted to some of the smaller islands as well
(e.g.. Heaney. 1986: Goodman & Ingle, 1993:
Heaney & Tabaranza. 1997; Musser et al., 1998;
Heaney & Mallari, 2002: Rickart et al.. 2002).
Evidence has suggested that those small islands
with endemic species are nearly always islands
surrounded by deep water (more than 125 m). so
that they were not connected to adjacent larger
islands during periods of low sea level during the
"ice ages" of the middle and late Pleistocene,
when sea level dropped to no more than 120 m
below present level (Heaney. 1986. 1991a.b:
Heaney & Regalado. 1998: Hanebuth et al.,
2000; Siddall e^l., 2003). Clearly, if we are to
understand the overall patterns of the evolution
and ecology of diversity in the Philippines, we
must understand the biodiversity patterns of
these smaller centers of endemism as well as the
larger ones.

One of the smaller Philippine islands sur-
rounded by deep water and still retaining
moderate rain-forest cover is Camiguin, located
about 10 km off the north-central shore of
Mindanao Island, with minimum intervening
water depth of 385 m (the island of Camiguin
Norte, which lies north of Luzon, is often
confused with Camiguin). It is a steeply moun-
tainous island (Fig. 1) of about 265 km2 with
several active volcanic cones that reach to
a maximum elevation of about 1620 m. A series
of biological surveys on Camiguin in the late
1960s (described below) that focused on birds
also yielded some mammal specimens. An earlier

report on the mammals (Heaney, 1984) conclud-
ed that the island had no endemic mammal
species and that it was depauperate relative to its
area. Subsequent studies on other islands made
us suspect that those earlier mammal surveys
were incomplete because so few mammal species
had been obtained and because the number of
nonvolant mammal voucher specimens was small
(thus indicating limited sampling effort). To
investigate the hypothesis that the previously
measured species richness of mammals was low
because of incomplete surveys, we returned to
Camiguin briefly in 1992 and more extensively in
1994 and 1995 to conduct additional mammal
inventories in all the major habitats along the
elevational gradient, especially by trapping small
mammals at higher elevations (Heaney & Taba-
ranza. 1997), and we obtained some new data on
the birds as well. In this volume, we summarize
results from both the 1960s and the 1990s
surveys and present evidence that both endemic
mammal and bird species are present on the
island (Heaney et al., 2006; Tello et al., 2006).

There is a second reason that we conducted
surveys on Camiguin in the mid-1990s. While the
Philippines is increasingly being recognized as
a global center for biodiversity, with unusually
high levels of endemism. it has simultaneously
vaulted into public view as one of the most
severely deforested of the tropical countries and
home to what may be the greatest concentration
of endangered species of mammals and birds
(Collar et al., 1994, 1999; Heaney & Regalado,
1998: Stattersfield et al., 1998; Mittermeier et al..

FIELDIANA: ZOOLOGY

■ 9°13

-9°11

124° 38'

9° 15'

124° 40'

124° 42'

124° 44'

124° 46'

124° 48'

White
Island

9°09

9 07'

9°05'

Mantigue

Island

o

5 kilometers

Fig. 2. Map of Camiguin Island showing the locations of mountain peaks referred to in the text, boundaries of
municipalities in the mid-1990s, the approximate boundaries of the proposed Timpoong-Hibok-hibok Natural
Monument (dotted line), and extent of forest cover in 1987 (gray area bounded by dashed line) from National
Mapping and Resource Information Authority (1988). The names of the municipalities are adjacent to the primary
population centers (= poblacion) of each municipality, shown as solid squares.

1999; Environmental Science for Social Change,
2000; Ong et al., 2002). Satellite maps of forest
cover from 1986 (National Mapping and Re-
source Information Authority, 1988) showed
a substantial area of forest cover in the center of
Camiguin (Fig. 2); such forest cover is now a rarity
in the Visayas (the islands of the central Philip-
pines) and adjacent regions (Heaney & Regalado,
1998; Environmental Science for Social Change,

2000), a portion of the country where the
concentration of endangered mammals and birds
is especially high (Wildlife Conservation Society
of the Philippines, 1997; Collar et al., 1999;
Heaney & Mallari, 2002). Because deforestation
has generally proceeded rapidly all over the
country in recent decades, we felt a sense of
urgency to obtain current information. Simulta-
neously, the prospect of finding species of

HEANEY AND TABARANZA: MAMMAL AND LAND BIRD STUDIES

- 9M5'

ST13'

124°42'

Site 9
ir

Site 1 1

Site 10

- gro9'

-gro7'

I I I I I I

5 kilometers

Fig. 3. Map of Camiguin Island showing 200-m elevational contours and locations of collecting sites in the
1990s (solid dots) and 1960s (dashed circles); as noted in the text, the locations of the latter are approximate. The
municipal population centers are shown also (solid squares).

mammals or birds in a place where enough forest
remained to support stable populations of any
endemic or endangered species was quite exciting,
posing the potential for positive action and the
development of successful conservation pro-
grams. We therefore proceeded to Camiguin with
a sense of cautious optimism and found condi-
tions that confirmed all our best hopes but also
some of our fears, as we shall describe here.

Geology, Vegetation, and Climate

Camiguin is composed almost entirely of
Quaternary volcanic material from currently
active volcanoes, and eruptions have occurred

within historic time (Hamilton. 1979; Mitchell et
al., 1986; Hall, 1998, 2002), with the most recent
serious eruptions in 1871 and 1951 (Agoo, 1995).
Five major and several secondary volcanic peaks
(Figs. 1-3) dominate the island's landscape and
serve as a tourist attraction; hikers often reach
the crater lake in Mt. Hibok-hibok and visit the
steam vents along its sides.

Camiguin is the northernmost, somewhat
isolated portion of the Central Mindanao Arc
of volcanic activity, which includes areas from
Mt. Apo in the south to Mts. Kalatungan and
Kitanglad in the center and Camiguin in the
north (Sajona et al., 1997). Volcanic activity in
this arc originated about 2.5 million years ago
(Ma), though most has taken place since about

FIELDIANA: ZOOLOGY

1.25 Ma. Camiguin is probably derived from
a single source of magma that first produced
undersea lavas in the late Pliocene or early
Pleistocene (ca. 2 Ma), with dryland appearing
not earlier than 1 Ma. Potassium-argon dating
shows the oldest dated magmatic materials on
the island to be those of the Mt. Ginsiliban and
Mt. Butay volcanoes in the southeastern corner
of the island (Fig. 1), which date to 0.34 Ma. The
peaks and volcanic flows associated with Mt.
Mambajao that form much of the center of the
island are much younger, and Mt. Hibok-hibok
in the northwestern portion of the island is the
youngest and only currently active vent (Pu-
nongbayan & Solidum, 1985; Sajona et al., 1997;
Castillo et al., 1999). We interpret this to indicate
that Camiguin originated as a dryland island not
earlier than 1 Ma and had high peaks and
substantial area by at least 300,000 years ago,
though possibly somewhat earlier (Heaney 1986,
1991a, b). The island is surrounded by deep
water, with a minimum depth to Mindanao of
385 m, which far exceeds the lowest documented
late or middle Pleistocene sea lowering of ca.
120 m (Siddall et al., 2003); thus, there is no
evidence of a dryland connection to Mindanao at
any time in the past.

Camiguin Island lies within a climate zone
characterized by annual rainfall of 2-3 m or
more in the lowlands, with seasonal variation
that includes a moderately dry period from
March to May (2.0-5.5 inches/mo; ca. 5-14 cm/
mo), especially wet from October to January
(10-15 inches/mo; ca. 25-38 cm/mo), and moist
during the rest of the year; mean annual rainfall
for a 24-year period at Mambajao was
99.4 inches (ca. 252.5 cm; Manalo, 1956). Tem-
perature declines and rainfall increases with an
increase in elevation; based on patterns elsewhere
in the Philippines (Heaney & Regalado, 1998),
we estimate that rainfall at 1500 m is roughly
triple that at sea level (i.e., about 7.5 m/year), but
no details from Camiguin are available.

The original vegetation of Camiguin was
lowland tropical rain forest from near the sea
to about 800 m elevation, with montane and
mossy forest to the peaks, but by the mid-1990s,
virtually no original vegetation remained below
300 m and little from 300 to about 800 m (Agoo,
1995). Beach and mangrove forest once occurred
along the coast, but little remains. Agoo (1995,
p. 4) described the upper elevations of Mt.
Hibok-hibok as a submontane plant community
"dominated by shrubby plants of Radermachera,

Rhododendron, Medinilla, and Vaccinium. Gym-
nosperms are conspicuously absent. . . . The
vegetation at the peak is devoid of tall trees
and shrubs. It is composed mostly of turfs of
grasses." Descriptions of vegetation at our
sampling areas during the 1990s are below.

Agoo (1995) noted the presence of the
following plants endemic to Camiguin: Miquelia
reticulata (Icacinaceae), Medinilla multiflora,
Memecylon subcaudatum (Melastomataceae), Sy-
zigium camiguense (Myrtaceae), Coelogyne con-
fusa, and Goodyera ramosii (Orchidaceae).

A small frog (Oreophryne nana) was the only
endemic vertebrate known from Camiguin when
we began our work (Alcala & Brown, 1998).

Methods

Prior Reports

Aside from a very small number of scattered
records (e.g., Gray 1843, who reported Paradox-
urus hermaphroditus) and a few specimens of birds
from the 1930s deposited at the Museum of
Comparative Zoology, the first significant surveys
of birds and mammals on Camiguin were
conducted by field teams from Silliman Univer-
sity at 10 sites in 1967, 1968, and 1969 and on the
small offshore island of Mantigue in 1969. The
field team in 1967 was led by D. P. Empesso and
R. B. Gonzales; we know of only bats collected by
this team, all of which were deposited in the Royal
Ontario Museum. Labels on specimens indicate
that collecting was conducted at three sites, listed
below as Sites 1 6—1 8. No field catalogs or notes
were kept, and no information is available on
their field procedures, but we suspect that they
followed roughly the same procedure as was used
in subsequent years by Dioscoro S. Rabor, as
follows. Field teams in 1968 and 1969, which
produced all the bird specimens and many of the
mammals cited here, were headed by Rabor; his
teams worked at seven sites, according to the
labels on specimens in the Field Museum of
Natural History (fmnh) and Delaware Museum
of Natural History (dmnh). B.R.T. Jr. worked on
several of Rabor's teams during the 1970s and
observed that Rabor followed a standardized
approach to fieldwork. The field crew simulta-
neously occupied a base camp, where Rabor
remained most of the time, and several satellite
camps. The satellite teams, made up of two or

HEANEY AND TABARANZA: MAMMAL AND LAND BIRD STUDIES

three persons, would go to areas chosen by Rabor
and operate for several days to several weeks.
Members in each satellite camp would set nets
each day. sometimes would run a small number of
large snap traps, and would hunt using shotguns
over an area that could readily be covered on foot.
Once each day. one member would hike to the
base camp with the specimens obtained since the
last such trip, then return to the satellite camp. A
similar team of collectors would operate in the
area of the base camp as well. At the time of
collection, specimens were labeled simply, usually
with the name of the site (often the name of
a barangay or sitio. which could cover many
square kilometers) and rough estimate of elevation
based on topographic maps. Rabor would work
with a team to prepare specimen labels (including
measurements, date, and locality data) and to skin
and stuff specimens. No field catalog or notes were
usually kept: data were recorded only on the
specimen skin tags (see also Rabor. 1966).

We have reconstructed the activities at Ra-
bor's field sites by listing all the locality names
that were noted and compiling the dates on
which specimens were obtained. Because field
teams operated simultaneously, the dates over-
lap. We noted the elevation, when present, for
each locality name, providing us with a range of
elevations for that site: many labels lack eleva-
tion, but we assume that the elevations noted
apply to all specimens from a given site. We
believe, however, that many of the elevations as
indicated on the labels of the specimens collected
during this period were overestimated and that
they should be used with caution. Additional
comments on elevational records can be found in
specific site descriptions.

Recent Data

Field studies that focused on mammals but
also obtained some data on birds were conducted
during three periods in 1992. 1994. and 1995 by
representatives from the fmnh and Mindanao
State University-Iligan Institute of Technology.
The first reconnaissance trip extended from 26 to
30 May 1992: bats were netted at a single
agricultural site by a field team led by A. T.
Peterson. The second period extended from 4
May 1994 to 4 June 1994: specimens were
collected at one agricultural site and three forest
sites by a team led by B.R.T. Jr. The third period
extended from 12 to 26 March 1995: specimens

were collected at two forest sites by a team led by
L.R.H. and B.R.T. Jr. Our methods for sampling
mammals and birds are presented in the relevant
chapters.

Study Sites

Camiguin is dominated by a series of volcanic
peaks, and most localities refer to those peaks.
We note that the three adjacent high peaks near
the center of the island (Fig. 3) are often
collectively referred to as Mt. Mambajao. though
we follow local people in referring to the
northeasternmost of the three major peaks as
Mt. Timpoong and the southwesternmost as Mt.
Mambajao.

Sites from 1990s

We surveyed mammals at one site in 1992.
four in 1994. and two in 1995: these are referred
to in the species accounts by number, and their
locations are shown in Figure 3. Additional
minor sites are described in the species accounts.
The dates include only those days when speci-
mens were obtained. Distances are given
from the municipal town centers, as shown in
Figure 2.

Site 1 — Barangay Balbagon. 1 km S. 2 km E
of Mambajao. 10 m. 9°14.5'N, 12444'E (28-29
May 1992). This site was situated in a highly
disturbed lowland agricultural area. The area
originally was lowland rain forest but in 1992
was a mosaic of agricultural fields, pastures, and
small bits of secondary lowland forest. Several
patches of shrubbery with heights of 2-4 m were
present, as were several large, thick patches of
erect bamboo. Six nets were set for a single night.
Bats and some birds were collected.

Site 2 — Barangay Balbagon. 7 km S. 2 km E
of Mambajao. 1000 m. 9r10.5'N. 124 44'E (28-
29 May 1992). A few specimens of birds were
netted in extensive, regenerating secondary forest
at this site.

Site 3 — Barangay Manuyog. Sagay Munici-
pality. 7 km S. 3 km W of Mahinog.' 0-300 m.
9°5.5'N, 124r45.5'E (5-11 May 1994). Sampling
was conducted in heavily disturbed lowland
agricultural land, with very few scattered patches
of degraded lowland forest along steep slopes
beside rivers. Trapping and netting were con-
ducted from close to the shoreline (ca. 50 m from
the beach) up to 300 m along the southwestern
slopes of Ginsiliban Peak, extending partially

FIELDIANA: ZOOLOGY

into Ginsiliban Municipality, for a total of three
net-nights and 148 trap-nights.

Site 4 — Barangay Kital-is, Sagay Municipali-
ty, Vi km N, 6Va km W of Mahinog, 1000 m,
9°9.5'N, 124°43.5'E (14-24 in May 1994). This
site was in disturbed transitional lowland/lower
montane forest along the southwestern slopes of
Mt. Mambajao at 900-1100 m elevation. The
slope was moderate, and the ground was covered
with abundant leaf litter. The vegetation con-
sisted mostly of small trees with dbh of 15-25 cm
and a few large trees standing 20-30 m high and
with 40-60 cm dbh; some lianas (5-10-cm di-
ameter), rattan {Calamus spp.), and climbing
bamboo {Schizostachyum spp.) clung to these
emergents. Ficus spp. were common. The most
common epiphytes were orchids, moss, and
ferns. Tree ferns (Cyathea spp.) and rattan
(Calamus spp.) seedlings were abundant as
ground cover. Scattered areas of humus were
up to 30 cm thick. Our total sampling effort at
this site consisted of 24 net-nights and 907 trap-
nights.

Site 5 — XA km N, 6 !/2 km W of Mahinog,
Sagay Municipality, 1200 m, 9°9.5'N, 124°43.5'E
(26-29 May 1994). This site was situated in
disturbed lower montane forest on moderate slope
at 1000-1300 m elevation, in the vicinity of Lasak-
lasak (Site 12). There was some evidence of small-
scale illegal logging done about a year previously.
Emergent trees had dbh of between 40 and 60 cm
and heights up to 30 m. Lianas and canopy
epiphytes (mainly ferns and mosses) were present.
Rattan {Calamus spp.) and climbing bamboo
{Schizostachyum spp.) were also present, with
pitcher plants {Nepenthes) present but rare. Un-
derstory and ground cover consisted of rattan
seedlings and ferns, with some sedges. Leaf litter
covered about 80% of the ground, and humus was
typically about 30 cm thick. Our total sampling
effort consisted of 339 trap-nights; at this site, nets
were concurrently maintained with those at Site 6
for a total of 14 net-nights.

Site 6 — Barangay Kital-is, Sagay Municipali-
ty, on a small peak near Mt. Mambajao, 1 km N,
7'/2km W of Mahinog, 1300m, 9°10'N,
124°43'E (26-29 May 1994). This site was
situated in primary mossy forest at 1200-
1400 m. Hanging moss was abundant on trees,
but moss cover on the ground was light and the
humus layer not more than a few centimeters
thick. Our total sampling effort consisted of 348
trap-nights for this site and 14 net-nights for
Sites 5 and 6 combined.

Site 7 — Mt. Timpoong, 2 km N, 6 Vi km W of
Mahinog, 1275 m, 9°10.5'N, 124°43.5'E (17-25
March 1995). This site was situated in primary
montane forest (Fig. 3) at 1225-1350 m eleva-
tion, in the vicinity of Lasak-lasak. The average
slope was ca. 35° and was often steep. The forest
had a fairly low and relatively open canopy; the
height of emergent trees was usually 20-25 m,
but a few reached 30 m; dbh was 12-30 cm, and
none had buttresses. The canopy was broken by
many treefalls, and canopy leaf sizes were small,
3-6 cm, often with serrated edges. Lower strata
leaf sizes were 4-20 cm and usually 6-10 cm.
Epiphytes were abundant, including ferns, or-
chids, and mosses. Arborescent pandans (Pan-
clanus sp.), melastome shrubs (Melastoma spp.),
and tree ferns (Cyathea spp.) were common, and
climbing rattan (Calamus spp.) and viney pan-
dans (Freycinetia spp.) were abundant; Ficus and
Musa were rare to absent. This area had thin to
moderate leaf litter cover and thin to moderate
humus (up to 15 cm deep); the soil consisted of
lightly weathered volcanic rock with many stones
at the surface and was generally very shallow.
There were no signs of human disturbance, but
many trees had fallen because of very shallow
soil, probably during occasional typhoons. There
were eight net-nights and 655 trap-nights.

Site 8— Mt. Timpoong, 2% km N, 6 '/a km W
of Mahinog, 1475 m, 9°11'N, 124°43.5'E (22-25
March 1995). Sampling at this site was con-
ducted in mossy forest. The site was on a steep
slope, averaging 50°, and ranged from 30-70°,
including steep gullies. The height of the canopy
was usually 8-10 m but varied from 2 to 3 m in
exposed spots to a maximum of ca. 18 m in low,
protected places; trees were generally gnarled,
with dbh of 12-20 cm (rarely 25 cm). Canopy
leaves were small, with serrated edges, typically
1-8 cm, but most were 4-5 cm. Oaks, laurels,
tree ferns, and arborescent Pandanus were
common. Lower-strata leaves were small, usually
4-8 cm. Ficus and Musa were rare to absent.
Epiphytes were abundant, including mosses,
ferns, orchids, and saplings, and pitcher plants
(Nepenthes spp.) were common. Canopy vines
(Freycinetia spp. and Calamus spp.) were abun-
dant. Ground plants included ferns, saplings,
and abundant pandans (Freycinetia). The ground
was covered by abundant and thick leaf litter
over deep humus (over a half meter thick, with
many tunnels and vacuities). Moss covered
nearly all tree trunks, branches, and fallen logs
and hung from branches in sheets and often

HEANEY AND TABARANZA: MAMMAL AND LAND BIRD STUDIES

covered the ground. No human disturbance was
seen, but there was evidence of some large
landslides and many tree falls. No netting was
performed at this site; there were 386 trap-
nights.

Sites from the 1960s

As noted above (see Methods), field teams
from Silliman University conducted surveys on
Camiguin at 10 sites in 1967-1969 plus an
additional site on the small, adjacent Mantigue
Island. As noted above (Methods), we have
estimated the location of these sites; our esti-
mates are shown in brackets. The vegetation
types and condition at these sites were not noted
by the collectors. We surmise, on the basis of
specimens collected here and the condition of the
remaining vegetation on Camiguin in the 1990s,
that when the collecting was done, the vegetation
below 800 m ranged from secondary lowland
forest to heavily populated agricultural areas. By
the time of our surveys in the 1990s, the lowland
forest was almost totally cleared and replaced
with coconuts, and hardly any area below 800 m
supported remnant forest except along steep
slopes. During the 1960s, from ca. 1000 m up to
the peak, the sites would have been covered with
primary montane and mossy forest, as they were
in the 1990s.

Site 9 — Mt. Catarman, Catarman Municipal-
ity, 2000-4950 ft (ca. 600-1500 m) [approx.
5.5 km S, 4.5 km W Mambajao, 9 12'N,
124 40.5' E] (10 29 June 1968). This was the
highest site surveyed in 1968; both birds and
mammals were collected. Mammals were collect-
ed at 2500-4500 ft (ca. 750-1400 m). The ma-
jority of the birds specimens were taken on 12-21
June, but a few others were collected on 10, 25,
and 29 June. Bird specimens were taken from
2000 to 4950 ft (ca. 600-1500 m). This would
suggest that the team sent to this camp concen-
trated their efforts from the middle to the upper
slopes, including the peak of Mt. Catarman, but
several were taken at 1000 ft (ca. 300 m) on 14
and 16 June 1968. However, we also know that
collecting at this site was done simultaneously
with Site 1 1 , and specimens from both sites were
then taken down to the base camp at Site 10 for
processing by D. S. Rabor. We suspect that in
the process, there was some mixing up of
elevational data for some specimens (see also
Site 1 1 ). Furthermore, the highest peak of the
Mt. Catarman does not exceed 1400 m, so all the

specimens cited by the team to have been
collected on this mountain above this elevation
were probably obtained close to the peak.

Site 10 — Gidag-on, Catarman Municipality,
500-1500 ft (ca. 1 50-450 m) [approx. 9T0.5'N,
124°39.5'E] (13-28 June 1968). This was the
lowest campsite during the 1968 field season;
birds and mammals were collected. Birds were
collected mostly from 23 to 28 June 1968, with
a few taken on earlier dates (13, 16-17, and 19
June). Some birds collected on 13 June were
noted to have been collected at 2000 ft (ca.
600 m), possibly the highest point reached from
this site. Because it was the lowest campsite in
1968, it is the most likely location of the base
camp that year. We are unable to locate this site,
but some collecting dates at this site overlapped
with Sites 9 and 1 1; we suspect that it was on the
lower slopes of Mt. Catarman. No information
on the vegetation when the specimens were
collected at this site is available, but we surmise
that it would have been partly secondary lowland
forest and partly agricultural.

Site 11 — Kasangsangan, Catarman Munici-
pality, 1000-2500 ft (300-800 m) [approx.
9 ll'N, 12440' E] (11-22 June 1968). This was
the middle campsite in 1968. We are unable to
locate this site, but because collecting dates
overlapped with Site 9, we suspect that it was
mainly on the lower to middle slopes of Mt.
Catarman. Several specimens were collected up
to 4950 ft (ca. 1500 m), which might indicate
that the team assigned to this site ventured all the
way to the peak of Mt. Catarman, but we suspect
mixing of localities, as noted above. No in-
formation on the vegetation when the specimens
were collected is available, but we surmise that,
similar to Site 9, it would have been partly
secondary lowland forest and partly agricultural.

Site 12 — Mt. Timpoong, Lasak-lasak, Mahi-
nog Municipality, 4400-5700 ft (ca. 1350-
1700m) [approx. 9 1 l'N, 12443. 5'E] (19-28
June 1969). We traveled through this site in
1995; it is located near the headwaters of the
Sagay River, from 1200 m to roughly the peak.
This was the highest site surveyed in 1969; birds
and mammals were collected. Collecting activi-
ties were conducted from 19 to 28 June. The
majority of the bird specimens were taken at
4800-5700 ft (ca. 1500-1700 m), indicating that
the team concentrated their collecting in the
high-elevation habitats on Mt. Timpoong. Sev-
eral specimens were taken slightly farther down,
at 4400 ft (ca. 1350 m). On 23 June 1969, some

FIELDIANA: ZOOLOGY

specimens attributed to this site were noted as
taken at 800 m. As with the previous year at Sites
9-11, collecting at this site was done simulta-
neously with Site 13, and specimens were taken
down to the base camp at Site 14 for processing.
We suspect that some mixing up of elevational
data for some specimens from among the three
sites occurred. While the vegetation at this site was
not indicated when the 1960s collecting was
undertaken, it would have been primary mossy
forest, based on the condition of the habitat that
we saw on Mt. Timpoong in the 1990s.

Site 13 — Mt. Timpoong, Matugnao, Mahinog
Municipality, 3150 ft(ca. 950 m) [approx. 9°10'N,
124°44.5'E] (12-26 June 1969). This was the
middle campsite established in 1969; birds and
mammals were collected. The majority of the bird
specimens bear the elevation 3150 ft (ca. 950 m),
with a few at 3250 ft (ca. 1000 m), around the
midslopes of Mt. Timpoong. A few specimens,
however, were noted as taken at 800 ft (ca. 250 m)
on 1 6 June and at 4800 ft (ca. 1 500 m) on 20 June.
Collecting at this site was done simultaneously
with Sites 12 and 14, and we suspect that some
mixing of elevational data occurred.

Site 14 — Puntod, Mahinog Municipality,
800 ft (ca. 250 m) [approx. 2 km N, 2 km W
Mahinog; 9°9.5'N, 124°46'E] (24-29 June 1969).
This was the lowest campsite established in 1969;
birds and mammals were collected. Bird collect-
ing took place on 24-29 June. Elevations were all
given as 800 ft (ca. 250 m), with a single bird
specimen bearing an elevation of 700 ft (ca.
200 m). Puntod is a village located on the lower
slopes of Mt. Timpoong, about 2 km west of the
town center of Mahinog (Fig. 3). This probably
was the base camp during the 1969 field season.
The vegetation in the 1960s is unknown, but
given its situation as a settled village during that
time, the habitat was probably already largely
agricultural with remnant lowland forest.

Site 15 — Mount Timpoong Peak, Mahinog
Municipality, 5700 ft (ca. 1600 m) [approx.
9°11'N, 124°43.5'E]. Only a single specimen of
Simcus murinus bears this locality. Given the
restriction to Mahinog Municipality, the site
must have been at the peak on the eastern side of
the mountain, near to or including Site 8. The
vegetation at this site would have been mossy
forest, as described for Site 8.

Site 16 — Mt. Mambajao, Mahidlaw, Catar-
man Municipality, 2500-3500 ft (ca. 800-
1000 m) [approx. 9 9.5'N, 124°42.5'E] (24 and
26 May 1967). Only a few bats were collected at

this site. Mt. Mambajao's highest peak is ca.
1600 m; this location in Catarman Municipality
suggests that specimens were taken from the
middle slopes, on the southern to southwestern
flank of the mountain.

Site 17 — Mt. Mambajao, Sangsangan, Catar-
man Municipality, 1400-3300 ft (ca. 400-
1000 m) [approx. 9°9'N, 124°42.5'E] (14-31
May, 15 June 1967). Only mammals were
collected at this site. This location suggests that
specimens were taken on the lower to middle
slopes of Mt. Mambajao, on the southern to
southwestern flank of the mountain.

Site 18 — Tag-ibo Cave, Catarman Municipal-
ity, 400 ft (ca. 100 m) (31 May 1967). A few bats
were collected at this site, which we have been
unable to locate. Because it was visited on the
same day as specimens were obtained from Site
17, they are probably near each other.

Site 19 — Mantigue Island [approx. 2 km N,
4 km E Mahinog; 9°10.5'N, 124°49.5'E] (28 June
1969). This site is a 4-ha coralline island, situated
ca. 3 km east of Barangay Hubangon, Mahinog
Municipality (Figs. 2 and 3). It was visited on 28
June 1969, and only White-collared Kingfisher
{Halcyon cloris), Yellow-vented Bulbul {Pycno-
notus goiavier), Pied Triller (Lalage nigra), and
Olive-backed Sunbird {Nectar una jugular is) were
collected. The vegetation when it was visited in
1969 was most likely beach forest, with some
disturbance.

Conservation

As documented by Balete et al. (2006), Heaney
et al. (2006), and Tello et al. (2006), Camiguin
supports at least 24 species of mammals and at
least 54 species of birds, and additional fieldwork
is likely to document the presence of additional
species. This includes two species of mammals (in
the genera Apomys and Bullimus) and one species
of bird (genus Loriculus) that are unique to
Camiguin; Camiguin is the smallest island in the
Philippines currently known to support unique
species of mammals (Heaney, 1986; Heaney et
al., 1998) and is smaller than any island pre-
viously known to support an endemic bird
(Peterson et al., 2000). The endemic species of
mammals have been documented to occur in
lowland, montane, and mossy forest from
1000 m to near the peaks but probably occur at
lower elevations as well where we were unable to
sample; the endemic bird (Tello et al., 2005) has
been documented from ca. 300 to 1200 m, which

HEANEY AND TABARANZA: MAMMAL AND LAND BIRD STUDIES

encompasses lowland and lower montane forest.
The presence of these unique species, along with
the frog and plants that are restricted to the
island, have caused Camiguin to be listed as a key
national and global priority site for conservation
(Mallari et al., 2001: Heaney & Mallari. 2002:
Ong et al.. 2002) that was overlooked by
previous assessments that were based on in-
complete biological surveys (e.g.. Hauge et al..
1986: Heaney. 1993: Peterson et al., 2000).

These diverse mammal and bird faunas
originally occurred along the entire elevational
and habitat gradient. The most fundamental
requirement for their conservation is the contin-
ued existence of substantial areas of all original
types of habitat in mature, good condition. The
areas must be large enough for the existence of
substantial populations, not just a few individu-
als, since populations often are vulnerable to
extinction when they drop below about 1000
individuals (Primack. 1998).

Fortunately. Camiguin retains enough forest
cover to make this possible, though there are
a number of challenges to surmount. The results
of the Swedish Satellite Survey of forest cover in
1987 (National Mapping and Resource Informa-
tion Authority. 1988) showed a fairly large area
of forest on Camiguin. mostly at upper eleva-
tions on Mt. Mambajao and Mt. Timpoong and
associated highlands, as shown in Figure 2. Our
surveys in 1994 and 1995 showed that much of
that forest still remained, though the edges had
been further degraded. Downslope. the degree of
disturbance progressively increased so that little
if any lowland forest remained in a primary
condition: indeed, most had been replaced with
coconut plantations, agricultural areas, and
grassland. In 2001. it was estimated that S29c
of the island, ca. 20.847 ha. was covered by
croplands and other inhabited areas: almost
half the island's area was covered with coco-
nut plantations (http://agrilO.norminet.org.ph/
NMProfile/profile_camiguin.htm). leaving \S9c
as secondary and primary forest.

According to local residents, much of the
forest on Camiguin was removed by commercial
logging operations in the 1980s, with the logged
areas subsequently being cleared for agriculture.
During the 1990s, small-scale commercial log-
ging, operating under salvage permits, was the
main agent responsible for the continuing de-
nudation of the areas below 1000 m. These
salvage permits technically allowed only dead
trees to be cut. but local residents told us that

trees were often killed deliberately by the loggers
so that they could then be cut down. Slash-and-
burn agriculture {kaingin) of the remnant forest
usually followed quickly (Heaney & Tabaranza.
1997). By the time we visited the island in 1995.
areas below 300 m had no native forest, and only
a little second growth remained up to ca. 800 m.
Only the montane and mossy forest was in
relatively good condition, but small-scale logging
and kaingin was creeping farther up on the
mountains: we noted a new kaingin at 1200 m on
the side of Mt. Timpoong in 1995 (Heaney &
Tabaranza. 1997: Mallari et al.. 2000).

It is essential to note that conservation of the
forest will have great benefits to the people of
Camiguin as well as to the wildlife. The forest
can serve as a permanent source of wood and
other nontimber forest products to the local
inhabitants if this is carefully managed. Perhaps
most crucially, the forests of Camiguin provide
the people with abundant water for domestic,
agricultural, and industrial uses. As an addition-
al essential ecological service, the forest helps
prevent soil erosion and landslides. In early
November 2002. about 200 human lives were lost
and thousands of houses and other properties
damaged by landslides during a typhoon, partic-
ularly in Mahinog. This disaster demonstrates
the great economic and human cost of forest
degradation. Additionally, the continued popular-
ity of Camiguin as a tourist destination will rely
strongly on its ability to provide beautiful scenery,
attractive areas for hiking, fresh and clean water,
and health} coral reefs (which require low levels of
siltation from adjacent rivers).

The importance of protecting the remaining
forest of Camiguin can be appreciated when one
considers its fast-growing population. In 1980, the
population of Camiguin was about 57.000; two
decades later, in 2000. the population had in-
creased to more than 73.000: in 2020. it is projected
to increase to 90.000 (http://www.popcom.gov.
ph/sppr/statistics/Ieastvis_wesmin_northmin.htm).
Consequently, the population density of Cami-
guin increased from 279 persons/km2 in 1980 to
306 persons/km" in 2000: these were well above
the national density of 160 persons/km2 and
251 persons/km2 in 1980 and 2000. respective-
ly (http://www.popcom.gov.ph/sppr/statistics/
lleastvisz_wesmin_northmin.htm: http://www.
popcorn. gov.ph/sppr/statistics/table3. htm).

In response to all these issues, including our
discoveries in 1994-1995 of endemic species of
mammals, the local gov ernment and Department

10

FIELDIANA: ZOOLOGY

of Environment and Natural Resources (DENR)
have moved steadily in recent years to have the
upland areas where forest remains declared the
"Timpoong-Hibok-hibok Natural Monument."
At the time of this writing, the process of
officially designating the protected area has
reached the office of the secretary of the DENR
and is expected to soon be endorsed to the
president. If it is successful at that level, as is
anticipated, it must then be voted on by the
Philippine Congress for final designation, a pro-
cess that may yet require several years. The
approximate proposed boundary of the natural
monument, with an area of 2,228 ha, plus
a 1,423-ha buffer zone, is shown in Figure 2.
Although both the location and area of forest
and the natural monument as shown in Figure 2
are approximate, it is clear that the majority of
the primary forest lies within the proposed
boundaries of the park.

We fully endorse these recommendations and
actions. But further, as part of this program of
environmental protection and stabilization, we
recommend the following to the local govern-
ment of Camiguin and the Philippine DENR as
essential activities: 1) continue and expand active
enforcement efforts to protect existing forests,
wildlife, and environment; 2) continue and
expand current reforestation projects, using only
native species of trees, not exotics, because the
exotic trees do not provide habitat for the
biodiversity of Camiguin and are injurious to
the soil; 3) cancel existing salvage cutting permits
for dead and fallen trees because these have
often been abused; and 4) complete the process
of declaring the remaining forest and key
watershed areas a national protected area as
soon as possible. To the maximum extent
possible, all portions of the protected area
should be connected by corridors of mature (or
regenerating) habitat, especially along rivers and
streams.

In connection with the above recommenda-
tions, it is essential to both wildlife conservation
and watershed protection that national and local
governments include substantial lowland areas
(those below 800 m) in the proposed national
park, regardless of the present condition of the
habitats and vegetation, especially including all
areas within 200 m of streams and rivers. We
therefore recommend that the natural monument
be expanded to include all good-quality second-
ary forest at all elevations and to include all
major watercourses down to at least 600 m of

elevation. This will enable future management to
rehabilitate/restore lowland forest, which is one
of the most critical habitats on Camiguin.

Acknowledgments

For assistance with the often challenging field
work on Camiguin, we thank N. Antoque, E.
Batara, N. Batocael, N. Bojo, M. Carmona, R.
Fernandez, A. DeOcampo, M. Jayoma, L.
Mostrales, A. T. Peterson, G. Rosell, A. Tabar-
anza, B. Tabaranza III, and D. Tabaranza.
Permits were provided by the Protected Areas
and Wildlife Bureau of the Department of
Environment and Natural Resources (DENR);
we offer special thanks to A. C. Alcala, C.
Custodio, M. Mendoza, and W. Pollisco and to
the Region 10 Office of the DENR. Emmauel
Aranas and Primitivo Espinas provided essential
assistance and logistical support on Camiguin
during the fieldwork. We thank E. Canete, C.
Custodio, and G. Rosell-Ambal for information
on the status of the protected area. For the loan of
specimens under their care, we are indebted to M.
Carleton and H. Kafka (usnm) and P. Myers
(ummz); we especially thank G. Hess (dmnh) for
much assistance and encouragement over many
years. The maps and graphs were prepared by
Lisa Kanellos; the photograph of Camiguin was
digitally prepared by Rebecca Banasiak. We
thank P. Janney for information on the geological
history of Camiguin and J. Bates, N. Collar, and
E. Rickart for constructive comments on an
earlier draft of the manuscript. This research
was supported by the World Environment and
Resources Program of the John D. and Catherine
T. MacArthur Foundation and by the Marshall
Field Fund, the Ellen Thorne Smith Fund, and
the Barbara Brown Fund for Mammal Research
of the Field Museum of Natural History.

Literature Cited

Agoo, M. G. 1995. Camiguin Island. Philippine
Biodiversity Information Center Plant Unit News-
letter, 3: 4.

Alcala, A. C, and W. C. Brown. 1998. Philippine
Amphibians: An Illustrated Guide. Bookmark,
Makati City.

Balete, D. S., B. R. Tabaranza, Jr., and L. R.
Heaney. 2006. An annotated checklist of the land
birds of Camiguin Island, Philippines, pp. 58-72. In

HEANEY AND TABARANZA: MAMMAL AND LAND BIRD STUDIES

11

Heaney, L. R., ed.. The Mammals and Birds of
Camiguin Island, Philippines, a Distinctive Center
of Biodiversity. Fieldiana Zoology, n. s., 106: 1-72.

Castillo, P. R., P. E. Janney, and R. U. Solidum. 1999.
Petrology and geochemistry of Camiguin Island,
southern Philippines: Insights to the source of adakites
and other lavas in a complex arc setting. Contributions
to Mineralogy and Petrology, 134: 33-51.

Collar, N. J., M. J. Crosby, and A. J. Stattersfield.
1994. Birds to Watch 2: The World List of
Threatened Birds. Birdlife International, Cam-
bridge.

Collar, N. J., N. A. Mallari, and B. R. Tabaranza,
Jr. 1999. Threatened Birds of the Philippines.
Bookmark, Makati City.

Dickinson. E. C. R. S. Kennedy, and K. C. Parkes.
1991. The Birds of the Philippines, an Annotated
Checklist. British Ornithologists" Union, Tring.

Environmental Science for Social Change. 2000.
Decline of the Philippine Forest. Bookmark. Ma-
nila. 44 pp. plus map.

Goodman. S. M., and N. R. Ingle. 1993. Sibuyan
Island in the Philippines — Threatened and in need of
conservation. Oryx. 27: 174-180.

Gray. J. E. 1843. List of the Specimens of Mammals in
the Collection of the British Museum. British
Museum (Natural History). London.

Hall. R. 1998. The plate tectonics of Cenozoic SE
Asia and the distribution of land and sea. pp. 99-
132. In Hall. R.. and J. D. Holloway. 1998.
Biogeography and Geological Evolution of SE Asia.
Backhuys, Leiden.

Hall. R. 2002. Cenozoic geological and plate tectonic
evolution of SE Asia and the SW Pacific: Computer-
based reconstructions, model and animations. Jour-
nal of Asian Earth Sciences. 20: 353-431.

Hamilton, W. 1979. Tectonics of the Indonesian
region. Geological Survey Professional Papers.
1078: 1-345.

Haneblth, T.. K. Stattegger, and P. M. Grootes.
2000. Rapid flooding of the Sunda Shelf: A late-
glacial sea-level record. Nature. 288: 1033 1035.

Hauge, P.. J. Terborgh, B. Winter, and J. Parkin-
son. 1986. Conservation priorities in the Philippines.
Forktail. 2: 83-91.

Heaney. L. R. 1984. Mammals from Camiguin Island.
Philippines. Proceedings of the Biological Society of
Washington. 97: 119-125.

— . 1986. Biogeography of mammals in Southeast
Asia: Estimates of rates of colonization, extinction,
and speciation. Biological Journal of the Linnean
Society. 28: 127-165.

— . 1991a. An analysis of patterns of distribution
and species richness among Philippine fruit bats
(Pteropodidae). Bulletin of the American Museum
of Natural History. 206: 145-167.

— . 1991b. A synopsis of climatic and vegetational
change in Southeast Asia. Climatic Change. 19: 53-61 .
. 1993. Biodiversity patterns and the conserva-
tion of mammals in the Philippines. Asia Life
Sciences. 2: 261-274.

Heaney, L. R., D. S. Balete, L. Dolar, A. C. Alcala,
A. Dans. P. C. Gonzales, N. Ingle, M. Lepiten, W.
Oliver, P. S. Ong, E. A. Rickart, B. R. Tabaranza,
Jr., and R. C. B. Utzurrum. 1998. A synopsis of the
mammalian fauna of the Philippine Islands. Fieldi-
ana Zoology, n.s.. 88: 1-61.

Heaney, L. R., and N. A. D. Mallari. 2002. A
preliminary analysis of current gaps in the pro-
tection of threatened Philippine terrestrial mammals.
Sylvatrop. 10(2000): 28-39.

Heaney, L. R.. and J. C. Regalado, Jr. 1998.

Vanishing Treasures of the Philippine Rain Forest.

The Field Museum. Chicago.
Heaney. L. R.. and B. R. Tabaranza. Jr. 1997. A

preliminary report on mammalian diversity and

conservation status of Camiauin Island, Philippines.

Sylvatrop, 5(1995): 57-64.

Heaney, L. R.. B. R. Tabaranza, Jr., D. S. Balete,
and N. Rigertas. 2006. Synopsis and biogeography
of the mammals of Camiguin Island. Philippines,
pp. 28^8. //; Heaney. L. R.. ed., The Mammals and
Birds of Camiguin Island. Philippines, a Distinctive
Center of Biodiversity. Fieldiana Zoology, n.s.. 106:
1-72.

Mallari. N. A. D. B. R. Tabaranza, Jr., and M. J.

Crosby. 2001. Key Conservation Sites in the

Philippines. Bookmark. Makati City.
Manalo, E. B. 1956. The distribution of rainfall in the

Philippines. Philippine Geographical Journal, 4:

104-166.

Mitchell, A. H. G.. F. Hernandez, and A. P. Dela
Cruz. 1986. Cenozoic evolution of the Philippine
Archipelago. Journal of Southeast Asian Earth
Sciences, 1: 3-22.

MlTTERMEIER, R. A., N. MYERS, P. RoBLES GlL, AND C.

G. Mittermeier. eds. 1999. Hotspots, Earth's
Biologically Richest and Most Endangered Terres-
trial Ecosystems. CEMEX. Mexico City.
Mittermeier. R. A., P. Robles Gil and C. G.
Mittermeier. eds.. 1997. Megadiversity: Earth's
Biologically Wealthiest Nations. CEMEX. Monter-
rey.

Mlsser. G. G.. L. R. Heaney. and B. R. Tabaranza,
Jr. 1998. Philippine rodents: Redefinition of
known species of Batomys (Muridae. Murinae) and
description of a new species from Dinagat Island.
Philippines. American Museum Novitates, 3237:
1-51.

National Mapping and Resource Information Au-
thority 1988. Sheet maps. 1 : 250,000. based on
satellite images from the Swedish Space Corpora-
tion. Manila. 53 sheets.

Ong. P.. L. E. Afiang. andR. G. Rosell-Ambal, eds.
2002. Philippine Biodiversity Conservation Priori-
ties: A Second Iteration of the National Biodiversity
Strategy and Action Plan. Philippine Department of
the Environment and Natural Resources. Quezon
City.

Peterson, A. T.. L. G. Ball, and K. W. Brady. 2000.
Distribution of the birds of the Philippines: Bio-
geography and conservation priorities. Bird Conser-
vation International. 10: 149-167.

12

FIELDIANA: ZOOLOGY

Primack, R. B. 1998. Essentials of Conservation
Biology. Sinauer Associates, Sunderland.

PUNONGBAYAN, R. S., AND R. U. SoLIDUM. 1985.

Camiguin Island. Philippine Institute of Volcanolo-

gy and Seismology, Quezon City.
Rabor, D. S. 1966. A report on the zoological

expeditions in the Philippines for the period 1961

1966. Silliman Journal, 13: 604-616.
Rickart, E. A., L. R. Heaney, and B. R. Tabaranza,

Jr. 2002. Review of Bullimus (Muridae: Murinae) and

description of a new species from Camiguin Island,

Philippines. Journal of Mammalogy, 83: 421-436.
Sajona, F. G., H. Bellon, R. C. Maury, M.

PUBELLIER, R. D. QUEBRAL, J. COTTEN, F. E. BAYON,

E. Pagado, and P. Pamatian. 1997. Tertiary and
Quaternary magmatism in Mindanao and Leyte
(Philippines): Geochronology, geochemistry, and
tectonic setting. Journal of Asian Earth Sciences,
15: 121-153.

SlDDALL, M., E. J. ROHLING, A. AlMOGI-LaBIN, C.
HEMLEBEN, D. MEISCHNER, I. SCHMELZER, AND D. A.

Smeed. 2003. Sea-level fluctuations during the last
glacial cycle. Nature, 423: 853-858.

Stattersfield, A. J. M., J. Crosby, A. J. Long, and
D. C. Wege. 1998. Endemic Bird Areas of the
World: Priorities for Conservation. Birdlife Interna-
tional, Cambridge.

Tello, J. G., J. F. Degner, J. M. Bates, and
D. E. Willard. 2006. A new species of Hang-
ing-Parrot (Aves: Psittacidae: Loriculus) from
Camiguin Island, Philippines, pp. 49-57. In Heaney,
L. R., ed., The Mammals and Birds of Cami-
guin Island, Philippines, a Distinctive Center of
Biodiversity. Fieldiana Zoology, n.s., 106: 1-
72.

Wildlife Conservation Society of the Philippines
1997. Philippine Red Data Book. Bookmark,
Makati.

HEANEY AND TABARANZA: MAMMAL AND LAND BIRD STUDIES

13

A New Species of Forest Mouse, Genus Apomys
(Mammalia: Rodentia: Muridae), from Camiguin
Island, Philippines

Lawrence R. Heaney1 and Bias R. Tabaranza, Jr.2

Abstract

An inventory of the mammals of Camiguin Island conducted in 1994 and 1995 documented
the presence of a previously unknown species of Philippine forest mouse of the endemic
Philippine genus Apomys, which is here named and described. Based on molecular data, the
new species is most closely related to two species {A. hylocoetes and A. insignis) from
Mindanao Island and to an unnamed species from Leyte, Biliran, and Bohol islands. The new
species is diagnosed in comparison to its three closest relatives on the basis of slightly browner
and less russet fur, slightly greater size overall, a moderately long and broad hind foot with
small plantar pads, large tail scales, slightly narrower zygomatic width and mastoid breadth,
deep rostrum of moderate length, a long orbit and braincase, narrow palate, large incisive
foramina, short distance from the posterior edge of the incisive foramina to the anterior edge
of the first upper molar, bony palate that extends well to the posterior of the posterior edge of
the last upper molar, bullae that are more strongly oriented toward the cranial midline axis,
third upper molar without a conspicuous anterolabial cusp, and a number of more subtle
features. It is one of two species of mammals now known to be endemic to Camiguin, the
other being Bullimus gamay (Rickart et al., 2002). Both are common in rain forest on
Camiguin Island at upper elevations. The presence of two endemic mammals on this small
(265 km") island is remarkable; there are no smaller islands in the Philippines known to
support endemic mammal species.

Introduction

The Philippine Islands are notable for their
large number of unique species of mammals; of
172 species known in 1998, 111 (64%) occurred
nowhere else in the world, one of the greatest

1 Field Museum of Natural History, 1400 South
Lake Shore Drive, Chicago, IL 60605-2496, U.S.A.

2 Department of Biology, Iligan Institute of Tech-
nology, Mindanao State University, Iligan City,
Lanao del Norte, Philippines.

concentrations of unique mammalian diversity
worldwide (Heaney et al., 1998). These species
are usually not widely distributed within the
Philippines but rather are confined to one or
a few islands. Recent studies have shown that the
geological history of the archipelago is largely
responsible for the pattern of distribution, with
most species of mammals found on only one of
the several islands that formed during periods of
low sea level in the late and middle Pleistocene.
Each of these Pleistocene islands is surrounded
by deep water (greater than 120 m current

14

FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, PP. 14-27

depth), and each has remained as an isolated
oceanic island throughout its existence. Howev-
er, though they are isolated by deep-water
channels, the channels are not wide, usually not
more than 25 km and often much narrower
(Heaney, 1986, 1991, 1993, 2004; Heaney and
Rickart, 1990; Heaney & Regalado, 1998).

Camiguin Island was noted by Heaney (1984,
1986) as an apparent exception to this pattern:
collections made on Camiguin in the 1960s by
field teams from Silliman University did not
include any endemic mammals, even though the
island seemed large enough to support them
(Heaney, 1986, 2004). After the discovery of four
endemic species on Sibuyan Island, another small
oceanic island in the archipelago (Goodman &
Ingle, 1993; Heaney et al., 1998), we suspected
that the mammals of Camiguin might not have
been fully surveyed, so we returned in 1994 and
1995 for further investigations (Heaney et al.,
2004; Heaney & Tabaranza, 2006). In the course
of those field studies, we documented the presence
of two previously unknown species of mammals
(Heaney & Tabaranza, 1997), Bullimus gamay
(Rickart et al., 2002) and a species of forest
mouse, genus Apomys. It is the purpose of this
paper to describe this new species of forest mouse.

Materials and Methods

Specimens examined for this study are housed
in the Delaware Museum of Natural History
(dmnh), Field Museum of Natural History
(fmnh), Mindanao State University-Iligan In-
stitute of Technology (msu-iit), National Muse-
um of the Philippines (nmp), University of
Michigan Museum of Zoology (ummz), and
United States National Museum of Natural
History (usnm). Half the specimens from Cami-
guin now housed in fmnh will be sent to nmp.
Material examined included specimens prepared
as study skins with skulls (and some with
postcranial skeletons), complete skeletons, and
formalin-fixed specimens stored in 70% ethyl
alcohol, many with skulls subsequently removed
and cleaned. The following samples were exam-
ined: Apomys hylocoetes — Mindanao Island:
Bukidnon Province: Mt. Kitanglad Range:
15 km S, 12.5 km W Dalwangan, elev. 2800 m,
fmnh 148055; 16.5 km S, 4 km E Camp Phillips,
elev. 1900 m, fmnh 147871-872, 148123-124;
18.5 km S, 4 km E camp Phillips, elev. 2250 m,

fmnh 147874-876, 147880, 147900-904, 147906,
148125-128, 148132, 148135-138. Apomys in-
signis — Mindanao Island: Bukidnon Province:
Mt. Kitanglad Range: 16.5 km S, 4 km E Camp
Phillips, elev. 1900 m, fmnh 148152; 17 km S,
7 km E Baungon, elev. 1550 m, fmnh 146703;
18 km S, 7 km E Baungon, elev. 1800 m, fmnh
146704-710, 146712-714, 146716-718, 147088-
089, 147091-092, 147094, 147098-099, 147102.
Apomys sp. — Biliran Island: 3 Vi km S, 5 Vi km
W Caibiran, elev. 700 m, ummz 160314, 160290,
160429^130; 4'/2 km S, 5 km W Caibiran, elev.,
920 m, ummz 160290-291, 160315-316. Leyte
Island: 9 km N, 3 km E Baybay, elev. 750 m,
ummz 160318; 8 !/2 km N, 2'/2 km E Baybay,
elev. 500 m, ummz 160317, 160441. Apomys n.
sp. — Holotype and referred specimens from
Camiguin Island (see below).

Specimens were assigned to age categories as
follows. Subadult animals are those that have
not completed cranial growth, especially those
having unfused basicranial sutures; these young
animals have pelage that is usually softer and
grayer than that of adults and are noticeably
lower in weight, and females are usually nullip-
arous. Young adults are older; they have unworn
adult pelage and have nearly completed cranial
growth but have not yet reached adult weight
and usually have not yet reproduced or are
pregnant for the first time. Adults have complet-
ed cranial growth and have adult pelage, and
usually the females are multiparous. Terminolo-
gy for description of external features follows
Brown (1971) and Brown and Yalden (1973).
Terminology for cranial and dental features
follows Musser and Heaney (1992). Scanning
electron micrographs of skulls and teeth were
made with uncoated specimens.

External measurements (total length, tail
length, hind foot length, length of ear from
notch, weight in grams) were taken from
collectors' field catalogs or specimen labels.
Fourteen cranial measurements (Table 1) were
taken with digital calipers to the nearest 0.01 mm
by Heaney; comparisons made in the text refer
only to specimens also measured by Heaney.

Results

The endemic Philippine genus Apomys was
described by Mearns (1905) to contain three
species: A. hylocoetes (as the type species), A.

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE

15

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16

FIELDIANA: ZOOLOGY

insignis, and A. petraeus (the last synonymized
with A. hylocoetes by Musser, 1982), all from Mt.
Apo, from which the name was derived. As
related by Musser (1982), additional species were
named from Luzon and Catanduanes, and the
genus was found throughout much of the
archipelago (Ruedas, 1995; Heaney et al.,
1998). However, because the initial description
was vague and many genera of Indo-Australian
rodents poorly studied, Apomys was briefly
synonymized with the genus Rattus at a time
when most Indo-Australian murids were placed
in that genus. Musser (1982) thoroughly rede-
scribed Apomys, pointing out its many distinctive
characters, and redefined the species then
known. Musser and Heaney (1992) further
defined and compared Apomys to other Philip-
pine murids, and they pointed out its apparent
close relationship to Chrotomys, Celaenomys,
and Rhynchomys, also endemic to the Philip-
pines. Using data from Musser and Heaney
(1992), Heaney and Rickart (1990) postulated
that Apomys was basal to the clade including
Chrotomys, Celaenomys, and Rhynchomys and
noted the diversification of this clade within the
Philippines as an example of adaptive radiation.
Heaney et al. (1998) noted the presence of many
undescribed species of Apomys, including the
species from Camiguin reported by Heaney and
Tabaranza (1997).

Rickart and Heaney (2002) showed that Ap-
omys hylocoetes, A. insignis, and the Leyte Apomys
(as well as most others from Greater Luzon and
Greater Negros-Panay) have distinctive karyo-
types, but the Apomys from Camiguin has not
been karyotyped. Steppan et al. (2003) used
molecular data to assess phylogenetic relation-
ships and geographic patterns of diversification
within Apomys. Analysis of variation in cyto-
chrome^ in 10 species (Fig. 1) showed the
presence of three major clades: one containing
Apomys datae and A. gracilirostris; a second clade
containing A. microdon, A. musculus, and two
undescribed species from Negros and Sibuyan (the
"Greater Luzon, Mindoro, and Negros clade");
and a third clade containing A. hylocoetes and A.
insignis from Mindanao, plus an undescribed
species from Leyte, Biliran, and Bohol and
another from Camiguin ( the "Greater Mindanao
clade"). The second and third of these three clades
form a monophyletic clade that Musser (1982)
described as being "species ... of small or
medium body size in which the canal for the
internal maxillary artery [also described as the

rC"

I A

A. "A/C" (Negros)
"B" (Sibuyan)

r

c

A. musculus
A. microdon
A. hylocoetes
A. insignis

A. "F" (Leyte)

A. camiguinensis

A. datae

A. gracilirostris

Rhynchomys isarogensis

Chrotomys gonzalesi

Fig. 1. Hypothesis of phylogenetic relationships
within the genus Apomys based on parsimony analysis
of cytochrome-b molecular data (based on Steppan et
al., 2003).

infraorbital branch of the stapedial artery] is
partially open and part of the artery is exposed
on the ventral surface of each pterygoid plate.
That feature, combined with bright tawny
upperparts, buffy underparts, brown or pale
buffy feet, and patterning on the tail (usually
monocolored or mottled, rarely sharply bi-
colored) suggest the five species [known in
1982] may be more closely related to each other
than to any others in Apomys." Musser's (1982)
recognition and diagnosis of this clade, based
on very few specimens, was prescient and
remains accurate. In the analysis by Steppan
et al. (2003), the species from Camiguin is basal
to the others in the "Greater Mindanao clade"
(Fig. 1), and they estimated the time of
divergence of the Camiguin mouse from the
other members of its clade at about 2.3 million
years ± about 25%.

We note that the Apomys from Biliran, Bohol,
and Leyte (which we hereafter refer to as "the
Leyte Apomys''') was tentatively considered to
represent either A. microdon (by Musser, 1982)
or A. littoralis (Rickart et al., 1993; Heaney et al.,

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE

17

Fig. 2. Photograph of an adult Apomys camiguinensis, taken on
March 1995.

Mt. Timpoong, Camiguin Island, in

1998). but the combination of the morphological
data presented here, karyotypic data (Rickart &
Heaney, 2002). and molecular data (Steppan et
al., 2003) have led us to conclude that the Leyte
Apomys is a distinct species; further details and
description will be published elsewhere.

The specimens from Camiguin are morpho-
logically similar to series of A. hylocoetes and A.
insignis from Mindanao and the undescribed
species from Leyte and Biliran but readily
distinguished from all three of these species on
the basis of external and cranial features. Heaney
et al. (1998) and Steppan et al. (2003) referred to
this animal informally as "'Apomys sp. D." We
now name the species from Camiguin as Apomys
camiguinensis, new species.

Apomys camiguinensis, new species

Holotype Adult male, fmnh 167878, collect-
ed 16 May 1994 by B. R. Tabaranza, Jr.
Specimen originally fixed in formalin, trans-
ferred to im ethyl alcohol with skull removed

and cleaned. Specimen is currently on deposit at
fmnh and is to be transferred to nmp.

Type Locality — Barangay Kital-is, Sagay Mu-
nicipality, 2 km N, 62 km W Mahinog, 1000 m
elevation, Camiguin Province, Camiguin Island.
9°9.5'N, 124°43.5'E (see Heaney & Tabaranza,
2006, for further details).

Referred Specimens and Localities — In addi-
tion to the holotype, 19 paratypes are known
from three localities ranging from 1000 to
1400 m (fmnh 154815-154816, 154854-154860,
167878-167882, plus 6 specimens at msu-iit); for
localities, see Heaney et al. (2006). All were
originally preserved in formalin and are now
stored in ethyl alcohol, many with skulls re-
moved and cleaned, or were prepared as
complete skeletons. Tissue samples are housed
at fmnh. Half of the specimens will be deposited
at nmp.

Distribution — Known only from the upper
elevations on Mt. Timpoong, Camiguin Island,
but probably occurring throughout the montane
and mossy rain forest on Camiguin Island (see
fig. 2 in Heaney et al., 2004) and possibly at
lower elevations.

IS

FIELDIANA: ZOOLOGY

Measurements — Table 1.

Etymology — The specific name refers to the
sole island on which the species is found. We
suggest the common English name "Camiguin
forest mouse."

Diagnosis — A species of the genus Apomys, as
defined by Musser (1982) and Musser and
Heaney (1992), including the following distinc-
tive generic features: rostrum long and moder-
ately narrow; viewed laterally, rostrum with
a rectangular shape, with premaxillaries project-
ing well anterior to the anterior edge of the
upper incisors; incisive foramina broad relative
to length; bony palate wide and long, densely
pitted and perforated; upper third molar re-
duced to a large round peg; lower third molar
also peg-like but retaining an anterior lamina,
without evidence of the two cusps that usually
form this lamina; occlusal surface of each first
and second upper molar consisting of two simple
chevron-shaped laminae followed by a small oval
lamina, without evidence of cuspidation; audito-
ry bulla separated from the squamosal and
alisphenoid by a gap that is formed by the
coalescence of the postglenoid foramen, the
postalar fissure, and the middle lacerate fora-
men.

As described in more detail in the following
section, the Camiguin mouse is defined by the
following characters or unique combination of
characters: moderate body size but somewhat
robust build for the genus overall; the tail is long
relative to body length, with unusually large
scales, and more often with a sharp transition
from dark brown dorsum to pale brown venter
(i.e., sharply bicolored) than in other species;
moderately long but unusually broad hind foot
with small plantar pads, and slightly shorter ear.
The pelage has more brown in the generally
russet-brown dorsum than on its closest relatives
and has more conspicuous salt-and-pepper
speckling dorsally; the mystacial and genal
vibrissae are long but moderate for the genus.
The cranium has an unusually long braincase
and orbit, somewhat narrow zygomatic and
mastoid width, and a moderately long but deep
and robust rostrum. The palate is rather narrow,
the posterior edge of the palate extends un-
usually far posterior to the last molar, the
incisive foramina long and wide, and the distance
from the posterior edge of the incisive foramina
is unusually short. The longest axis of the bullae
is about 35° from the midline axis of the skull.
The toothrows are of moderate size; the ante-

rolabial cusp of the third upper molar is barely
evident in most individuals.

Description and Comparisons — Apomys cami-
guinensis is an attractive mouse with large eyes
and ears, long tail, and soft pelage (see Frontis-
piece, this volume, and Fig. 2). As with other
members of the genus, the pelage is soft and
dense, without spines or stiff hairs. The dorsal
coloration is a rich brownish-russet with a small
amount of salt-and-pepper speckling; underfur is
pale slate-gray. The venter is paler, usually
nearly white with a wash of buffy or pale russet,
but some individuals have blazes of pure white
(usually on the chest) or are much darker brown
or russet-brown. There is a narrow area of bare
skin around the eyes; the ears are moderately
dark brown, with short hairs apparent on the
outer surface, and present but tiny and nearly
invisible on the inner surface. The mystacial
vibrissae are long and conspicuous. The dorsal
surface of the fore and hind feet are mostly buffy
or pale brown but with a narrow band of
scattered darker hairs around the midline, and
these decrease in number and length toward the
distal end of each foot (with only pale hairs on
the dorsal surface of the toes). The feet are lightly
pigmented or unpigmented on the ventral
surface, with conspicuous plantar pads on the
ventral surface (Fig. 3). The tail is long with
conspicuous scales; fine hairs that are present
between the scales are most visible on the dorsal
and lateral surfaces and least visible ventrally.
The tail is darker on the dorsal surface than on
the ventral surface. The scrotum of adult males is
fairly small and projects beyond the abdomen
only partially on the posterior portion and has
black or dark brown pigment at the posterior tip,
about 3-5 mm in length. Females have two pairs
of inguinal mammae.

Apomys camiguinensis is easily distinguished
from most members of the genus by its in-
termediate size (only A. insignis and A. hylo-
coetes are similar) and from those two species by
both external and cranial characters. Apomys
camiguinensis has total length (average 254—
260 mm) slightly greater than A. hylocoetes
(248-252 mm) and A. insignis (251-252 mm)
and substantially greater than the Leyte Apomys
(238-245 mm; Table 1). The average tail length
(146-148) is equal to that of A. insignis (147—
148 mm), and substantially longer than that of
A. hylocoetes (141-142 mm) and the Leyte
Apomys (140-145 mm). The tail averages 57%
of the total length in A. camiguinensis, compared

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE

19

Fig. 3. Ventral surface of the right hind feet of (A) Apomys camiguinensis, (B) A. hylocoetes, (C) A. insignis,
and (D) the undescribed Apomys from Leyte, all to roughly the same scale. Those of A. hylocoetes and A. insignis
are redrawn from Musser (1982).

to 56.5% in A. hylocoetes, 58% in A. insignis, and
59% in the Leyte Apomys. The hind foot (about
33 mm) is about equal in length to that of A.
insignis (33 mm) and is substantially longer than
in A. hylocoetes (31-32 mm) and the Leyte
Apomys (30-31 mm); the hind foot of A.
hylocoetes is notably the broadest (Fig. 3; see
also fig. 7 in Musser, 1982). Ear length is greatest
in A. hylocoetes (20.2 mm), with A. insignis
(19.5 mm), A. camiguinensis (19.0 mm), and the
Leyte Apomys (18-19 mm) progressively slightly
smaller (Table 1 ). The weight of A. camiguinensis
(38-41 g) averages the greatest of the four,
followed by A. hylocoetes (36-39 g), A. insignis
(37 38 g), and the Leyte Apomys (28-31 g). In
other words, the Camiguin Apomys is relatively
heavy and long and has a relatively long tail (but
slightly less long proportionately than some close
relatives), a moderately long hind foot, and
somewhat short ear.

The following qualitative external characters
also distinguish Apomys camiguinensis from its
three closest relatives. The dorsal coloration of
the Camiguin mouse is less russet and more
brown than in A. hylocoetes and A. insignis, with

more of the salt-and-pepper appearance; the
Leyte mouse is dorsally brighter red than the
others, with more red and orange in the russet
than the other three and almost no salt-and-
pepper. Ventral coloration is generally similar in
all four, though with the variation noted above,
but A. hylocoetes tends to have more of an
orange wash than the others. The dorsal surface
of the hind feet usually is palest in the Leyte
mouse and darkest in the Camiguin mouse. The
mystacial vibrissae are long on all four species
but longest on A. insignis (up to 56-60 mm
maximum), on which they reach past the middle
of the back, and intermediate on the Camiguin
mouse (52-55 mm), A. hylocoetes (51-55 mm),
and Leyte mouse (50-55 mm). Genal vibrissae
reach to the anterior edge of the largest lateral
pad on the Camiguin and Leyte mice but farther
forward, to the base of the toes, on A. hylocoetes
and A. insignis.

The hind foot (Fig. 3) differs markedly among
the four. The foot of A. camiguinensis is about
the same length as that of A. hylocoetes but is
broader and has smaller plantar pads. The hind
foot of the Leyte mouse is proportioned similarly

20

FIELDIANA: ZOOLOGY

Fig. 4. Photographs of dorsal, ventral, and lateral views of the crania of Apomys camiguinensis (A; fmnh
167878, holotype), A. hylocoetes (B; fmnh 148146), A. insignis (C; fmnh 147092), and the undescribed Apomys
from Leyte (D; ummz 160290), all to the same scale.

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE

21

Fig. 5. Scanning electron micrographs of the ventral view of the anterior portion of the skulls of Apomys
eamiguinensis (A; fmnh 167878, holotype), A. hylocoetes (B; fmnh 148146), A. insignis (C; fmnh 147092), and the
undescribed Apomys from Leyte (D; ummz 160290), all to same scale.

to A. hylocoetes but is smaller. The hind foot of sturdy and sharply pointed; those of A. hylo-

A. insignis is very long and narrow, with plantar
pads of moderate size. On all species, the claws
on all five digits are unpigmented and laterally
compressed. On A. eamiguinensis, the claws are

eoetes are slightly thinner and more sharply
pointed and the digits bearing them slightly more
slender. The digits and claws of A. insignis are
shorter and more slender; the digits and claws of

FIELDIANA: ZOOLOGY

Fig. 6. Scanning electron micrographs of the ventral view of the posterior portion of the skulls of Apomys
camiguinensis (A; fmnh 167878, holotype), A. hylocoetes (B; fmnh 148146), A. insignis (C; fmnh 147092), and the
undescribed Apomys from Leyte (D; ummz 160290), all to same scale.

the Leyte mouse are still smaller and more
slender. The forefeet of A. camiguinensis have
smaller pads than those of the others (but only
proportionately in the case of the generally
smaller Leyte mouse), but the feet are slightly
broader and more robust than in the other
species. All four species have a flat, unpigmented
nail on the pollex. The relative size and thickness
follows the same pattern as the hind feet, with A.

camiguinensis being the most robust, A. hylo-
coetes slightly shorter and thinner, A. insignis
much shorter and more slender, and the Leyte
mouse much like A. insignis but a bit smaller
overall.

The tail scales on the Camiguin mouse are
largest, with 12-12.5 scales/cm near the base,
compared to 14-15 scales/cm in A. hylocoetes,
13-14 in A. insignis, and 13-15 in the Leyte

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE

Fig 7 Scanning electron micrographs of the occlusal surface of the maxillary toothrows of Apomys
camiguinensis (A; fmnh 167878, holotype), A. hylocoetes (B; fmnh 148146), A. insignis (C; fmnh 147092), and the
undescribed Apomys from Leyte (D; ummz 160290), all to same scale.

Fig. 8. Scanning electron micrographs of the occlusal surface of the mandibular molariform toothrows of
Apomys camiguinensis (A; fmnh 167878, holotype), A. hylocoetes (B; fmnh 148146), A. insignis (C; fmnh 147092),
and the undescribed Apomys from Leyte (D; ummz 160290), all to same scale.

24

FIELDIANA: ZOOLOGY

mouse. The size of the scales remains about the
same more distally on a given species to about
the midpoint; from there to the tip scale size
decreases, mostly in the last quarter of the
length, to about half the length (and one-fourth
the size) of scales near the base of the tail. The
scales of the tail are dark brown dorsally and
nearly white ventrally on all four species, but the
transition is most often an abrupt line on the
Camiguin mouse, often producing a sharply
bicolored tail, and most often gradual on the
other three species. Typically, three hairs grow
from beneath each scale, projecting only slightly
laterally from the tail; on the basal quarter of the
tail, one of the three hairs is missing from a given
scale about one-third of the time. On A.
camiguinensis, the hairs are about one and one-
third the length of a scale near the base of the
tail, have about the same length 25% toward the
tip, are slightly longer at the midpoint, are about
one and two-thirds the length of a scale 75%
toward the tip, and are about five times the
length of a scale near the tip, where scale
length has been reduced by about half relative
to the base. On A. hylocoetes, hairs are about
one and a half the length of a scale near the base
of the tail, are similar 25% toward the tip, are
about twice the length of a scale near the
midpoint, are about two and a half times
the length of a scale 75% toward the tip, and
are more than five times the length of a scale
near the tip. On A. insignis, hairs are one and
one-third a scale length near the tail's base,
similar at 25%, one and two-thirds near the
midpoint, about two and one-half of a scale
length near 75%, and about three and one-half
the length of a scale near the tip. On the Leyte
mouse, tail hairs are slightly less than the
length of a scale near the base of the tail, similar
at the midpoint and at 75%, and about two times
the length of a scale near the tip. In all four
species, the hair becomes slightly greater in
diameter distally than it is proximally; combined
with the trend for greater length, this means
that the tail become more heavily covered with
hair distally. None of the species shows elongat-
ed hairs at the very tip of the tail (i.e., there is
no "pencilling"). Hairs on the dorsal surface
are more heavily pigmented dorsally than
ventrally in all four species. On adults of all four
species, the dorsal surface of the tip (2-5 mm) of
the tail becomes worn, with most scales and
many hairs absent, leaving a smooth, leathery
surface.

The cranium of Apomys camiguinensis (Figs. 4-
6) has basioccipital length (28.3-29.0 mm) slight-
ly greater than that of A. hylocoetes (28.3-
28.8 mm), clearly more than A. insignis (27.8-
28.0 mm) and substantially more than the Leyte
Apomys (25.3-26.3 mm); the interorbital width is
proportioned similarly (Table 1). Zygomatic
breadth averages greatest in A. hylocoetes
(15.0-15.2 mm), followed by A. camiguinensis
(14.9-15.0 mm), A. insignis (14.7-14.8 mm), and
the Leyte Apomys (13.1-13.7 mm), with mastoid
breadth following the same pattern (Table 1).
Nasal length averages substantially greater
in A. hylocoetes (12.3-12.5 mm) than in A.
insignis (11.6-11.9 mm), A. camiguinensis (11.1-
11.4 mm), or the Leyte Apomys (10.3-10.7 mm),
and anterior nasal breadth is similarly patterned
(Table 1). Rostral depth averages greatest in A.
camiguinensis (6.3-6.4 mm), with A. hylocoetes
(6.2-6.3 mm) and A. insignis (6.3 mm) slightly
less deep and very similar to each other and the
Leyte Apomys generally less deep (5.9-6.1 mm).
Rostral length (Fig. 5) is clearly greatest in A.
hylocoetes (13.1-13.2 mm), with A. camiguinensis
(11.6-12.0 mm) and A. insignis (11.9-12.0 mm)
similar to each other but much greater than the
Leyte Apomys (10.6-11.2 mm). Orbital length
averages greatest in A. camiguinensis (10.2-
10.3 mm), followed by A. insignis (10.0-
10.2 mm) and A. hylocoetes (9.7-10.0) and the
small Leyte Apomys (9.2-9.6). To summarize, A.
camiguinensis is characterized by the longest
skull and deepest rostrum, but A. hylocoetes
has slightly greater zygomatic and mastoid width
as well as greater nasal length and breadth and
rostral length. Although the rostrum of A.
camiguinensis is moderate in length, the orbital
region and braincase are unusually long. The
cranium of A. insignis tends to be smaller but
generally similarly proportioned to A. camigui-
nensis, and the Leyte Apomys is smaller in all
dimensions, though it seems to follow the pattern
of A. camiguinensis and A. insignis.

The maxillary toothrow (Fig. 7) of A. cami-
guinensis (6.0 mm) averages nearly identical in
length to that of A. hylocoetes (6.0 mm) and A.
insignis (6.0 mm), and all are much greater than
that of the Leyte Apomys (5.2-5.3 mm). Palatal
breadth at M1 (Fig. 4) is greatest in A. insignis
(6.5-6.6 mm), followed by A. hylocoetes
(6.5 mm) and A. camiguinensis (6.3-6.5 mm)
and the Leyte Apomys (5.7-5.8 mm). Diastema
length (Figs. 4 and 5) in A. camiguinensis
(7.3 mm) averages slightly greater than in A.

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE

25

hvlocoetes (7.2 -7.3 mm), more than A. insignis
(7.0 mm), and least in the Leyte Apomys (6.5
6.8 mm). In other words, the maxillary tooth-
rows of the three larger Apomys are all similar,
with the Leyte Apomys having disproportionate-
ly short toothrow, but the palate of A. camigui-
ncnsis is disproportionately narrow (as is that of
the Leyte Apomys), and that of A. hvlocoetes is
disproportionately wide.

In addition, we note the following qualitative
characters. The incisive foramina (Fig. 5) are
widest in A. eamiguinensis and in A. hvlocoetes
and longest in A. eamiguinensis and A. insignis, so
that the area of the foramina is greatest in A.
eamiguinensis. The distance from the posterior
edge of the incisive foramina to a line between the
anterior edges of the first maxillary molar is
shortest in A. eamiguinensis, slightly greater in
A. insignis, and longest in A. hvlocoetes and
the Apomys from Leyte. The braincase of A.
eamiguinensis is slightly more elongate and that of
the Leyte Apomys proportionately most squarish
among the four species (Fig. 6). The posterior
edge of the bony palate (Fig. 6) extends farthest
posterior to the posterior edge of the last
maxillary molar in A. eamiguinensis, slightly less
far in A. hvlocoetes and the Apomys from Leyte.
and least far in A. insignis. The long axis of the
bullae (Fig. 6) is about 45 from the cranial
midline axis in the Apomys from Leyte, about 40
in A. hvlocoetes and A. insignis, and about 35c in
A. eamiguinensis. The hard palate of A. hvlocoetes
and the Leyte Apomys are usually most heavily
pitted and perforated with vacuities and A.
eamiguinensis and A. insignis less so (Figs. 4 and
7). The third upper molar (Fig. 7) has an
anterolabial cusp (probably tl; see Musser &
Heaney, 1992, p. 65) that is well developed in A.
hvlocoetes, less conspicuous in A. insignis, and
barely evident in A. eamiguinensis and the Apomys
from Leyte. The first upper molar of A. hvlocoetes
tends to have a more conspicuous anterolingual
cleft than do the other three species (Fig. 7). Both
upper and lower toothrows of A. insignis are the
most massive (Figs. 7 and 8), with A. eamigui-
nensis somewhat less massive, A. hvlocoetes sub-
stantially less so, and the Apomys from Leyte
smallest overall. All four species have the canal
for the infraorbital branch of the stapedial artery
partially open and part of the artery exposed on
the ventral surface of each pterygoid plate (Fig. 6),
as noted by Musser (1982).

Ecology Sec Heaney et al. (2006) for ecolog-
ical information

Discussion

The presence of Apomys eamiguinensis as an
endemic species on a small island, along with the
additional murine rodent Bullimus gamay (Rick-
art et al., 2002) and the Hanging-Parrot (Lor-
iculus sp., described in this volume), as noted
above, clearly indicates the importance of
Camiguin Island as a unique center of biological
diversity that is worthy and in need of conser-
vation (Heaney & Tabaranza, 2005). In addition,
the distinctiveness of this species confirms pre-
dictions made on the basis of biogeographic
models (Heaney, 2004) of the expected presence
of endemic small mammals on Camiguin. Fur-
ther studies of the mammals, birds, and other
organisms are clearly warranted to determine,
for example, the degree of genetic difference
from closest relatives (most of which occur on
Mindanao) as a means of assessing the role of
colonization and gene flow in determining
patterns of species richness and endemism in
the Philippines. In other words, Camiguin
represents a natural experiment, as a young
oceanic, volcanic island that is near to a large,
rich source of species (Mindanao), in which we
can measure the impact of genetic isolation in
animals and plants of varying vagility under
standardized conditions. Such studies are certain
to produce new insights into the process by
which biological diversity is generated in the
Philippines and in other oceanic archipelagos.

Acknowledgments

We thank Rebecca Banasiak for assistance in
photographing skulls, Betty Strack for assistance
with making scanning electron microscope
images, and Lisa Kanellos for preparation of
figures; Figure 3D was drawn by Jodi Sedlock.
The Protected Areas and Wildlife Bureau of the
Philippine Department of Environment and
Natural Resources (denr) provided encourage-
ment and permits, and the Region 10 office of
the denr provided logistical support. Specimens
for comparison were kindly loaned by G. Hess
(dmnh), P. Myers (ummz), and M. Carleton, L.
Gordon, and H. Kafka (usnm). The manuscript
was much improved by comments from D. S.
Balete, G. G. Musser, P. Myers, and E. A.
Rickart. Funding was provided by the Marshall
Field Fund, Ellen Thorne Smith Fund, and the

26

FIELDIANA: ZOOLOGY

Barbara Brown Fund for Mammal Research of
the Field Museum of Natural History.

Literature Cited

Brown, J. C. 1971. The description of mammals, 1.
The external characters of the head. Mammal
Review, 1: 151-168.

Brown, J. C, and D. W. Yalden. 1973. The
description of mammals, 2. Limbs and locomotion
of terrestrial mammals. Mammal Review, 3:
107-134.

Goodman, S. M., and N. R. Ingle. 1993. Sibuyan

Island in the Philippines — Threatened and in need of

conservation. Oryx, 27: 174-180.
Heaney, L. R. 1984. Mammals from Camiguin Island,

Philippines. Proceedings of the Biological Society of

Washington, 97: 119-125.

. 1986. Biogeography of mammals in Southeast

Asia: Estimates of rates of colonization, extinction,
and speciation. Biological Journal of the Linnean
Society, 28: 127-165.

1991. An analysis of patterns of distribution

and species richness among Philippine fruit bats
(Pteropodidae). Bulletin of the American Museum
of Natural History, 206: 145-167.

-. 1993. Biodiversity patterns and the conserva-

tion of mammals in the Philippines. Asia Life
Sciences, 2: 261-274.

2004. Conservation biogeography in oceanic

archipelagoes, pp. 345-360. In Lomolino, M. V. and
L. R. Heaney, eds., Frontiers of Biogeography: New
Directions in the Geography of Nature. Sinauer
Associates, Sunderland.

Heaney, L. R., D. S. Balete, L. Dolar, A. C.
Alcala, A. Dans, P. C. Gonzales, N. R. Ingle, M.
Lepiten, P. S. Ong, W. L. R. Oliver, E. A. Rickart,
B. R. Tabaranza, Jr., and R. C. B. Utzurrum.
1998. A synopsis of the mammalian fauna of the
Philippine Islands. Fieldiana Zoology, n.s., 88: 1-
61.

Heaney, L. R., and J. C. Regalado, Jr. 1998.
Vanishing Treasures of the Philippine Rain Forest.
The Field Museum, Chicago.

Heaney, L. R., and E. A. Rickart. 1990. Correlations
of clades and clines: Geographic, elevational, and
phylogenetic distribution patterns among Philippine
mammals, pp. 321-332. In Peters, G. and R.
Hutterer, eds., Vertebrates in the Tropics. Museum
Alexander Koenig, Bonn.

Heaney, L. R., and B. R. Tabaranza, Jr. 1997. A
preliminary report on mammalian diversity and
conservation status of Camiguin Island, Philippines.
Sylvatrop, 5(1995): 57-64.

. 2006. Introduction to the mammal and land

bird faunas of Camiguin Island, Philippines, pp. 1-
13. In Heaney, L. R., ed., The Mammals and Birds of
Camiguin Island, Philippines, a Distinctive Center of
Biodiversity. Fieldiana Zoology, n.s., 106: 1-72.

Heaney, L. R., B. R. Tabaranza, Jr., D. S. Balete,
and N. Rigertas. 2006. Synopsis and biogeography
of the mammals of Camiguin Island, Philippines,
pp. 00-00. In Heaney, R. L., ed., The Mammals and
Birds of Camiguin Island, Philippines, a Distinctive
Center of Biodiversity. Fieldiana Zoology, n.s., 106:
28-48.

Mearns, E. A. 1905. Descriptions of new genera and
species of mammals from the Philippine Islands.
Proceedings of the United States National Museum
of Natural History, 28: 425-460.

Musser, G. G. 1982. Results of the Archbold
Expeditions, No. 108. The definition of Apomys,
a native rat of the Philippine Islands. American
Museum Novitates, 2746: 1-43.

Musser, G. G., and L. R. Heaney. 1992. Philippine
rodents: Definitions of Tarsomys and Limnomys
plus a preliminary assessment of phylogenetic
patterns among native Philippine murines (Murinae,
Muridae). Bulletin of the American Museum of
Natural History, 211: 1-138.

Rickart, E. A., and L. R. Heaney. 2002. Further
studies on the chromosomes of Philippine rodents
(Muridae: Murinae). Proceedings of the Biological
Society of Washington, 115: 473^87.

Rickart, E. A., L. R. Heaney, P. D. Heideman, and
R. C. B. Utzurrum. 1993. The distribution and
ecology of mammals on Leyte, Biliran, and Maripipi
Islands, Philippines. Fieldiana Zoology, n.s., 72:
1-62.

Rickart, E. A., L. R. Heaney, and B. R. Tabaranza,
Jr. 2002. Review of Bullimus (Muridae: Murinae)
and description of a new species from Camiguin
Island, Philippines. Journal of Mammalogy, 83:
421^136.

Ruedas, L. A. 1995. Description of a new large-bodied
species of Apomys Mearns 1905 (Mammalia:
Rodentia: Muridae) from Mindoro Island, Philip-
pines. Proceedings of the Biological Society of
Washington, 108: 302-318.

Steppan, S., C. Zawadski, and L. R. Heaney. 2003. A
molecular assessment of phylogenetic relationships
and patterns of phylogenesis in the Philippine murid
rodent Apomys. Biological Journal of the Linnean
Society, 80: 699-715.

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE

27

Synopsis and Biogeography of the Mammals of Camiguin
Island, Philippines

Lawrence R. Heaney,1 Bias R. Tabaranza, Jr.,1 Danilo S. Balete,1,3 and
Natalie Rigertas1

Abstract

Biodiversity surveys in the 1960s and 1990s on Camiguin Island, a geologically young,
volcanically active oceanic island surrounded by deep water, have demonstrated the presence
of 24 species of land mammals. Three species (one insectivore and two rodents) are not native
to the Philippines, but all others (one insectivore, 12 bats, one monkey, four rodents, two
small carnivores, and one ungulate) are indigenous. Among those captured in the 1990s were
two previously unknown species of murid rodents in the genera Apomys and Bullimus that are
endemic to Camiguin. The discovery of two new species on such a small island (ca. 265 km ) is
remarkable; Camiguin is currently the smallest island in the Philippines known to have unique
species of mammals. Total species richness of nonvolant mammals on Camiguin is low, but
relative to islands that were once part of Pleistocene Greater Mindanao, Camiguin is not
depauperate. However, its fauna is not ecologically balanced in the same way as the faunas of
the islands that were part of Greater Mindanao: ground-living shrews (Crocidura) and rodents
(Apomys, Bullimus, Crunomys, and Rat t us) from lowland forest, along with some large
mammals (Macaco, Paratloxwus, and Sus) are well represented on Camiguin, but all the
arboreal small mammals that characterize lowland forest on Mindanao (Sundasciurus,
Exilisciurus, Cynocephalus, and Tarsius), ground-living small mammals from montane
habitats ( Urogale, Podogymnura, Batomys, Limnomys, and Tarsomys), and one large mammal
(Cervus) are absent. Additionally, at least two genera of fruit bats (Haplonycteris and
Megaerops) that are fairly common in lowland rain forests on Mindanao are absent on
Camiguin. The presence of some nonvolant mammals demonstrates that dispersal across the
deep but narrow intervening channel takes place, but the presence of two species endemic to
Camiguin and the absence of other species that are present on nearby Mindanao implies that
dispersal probably is rare. The Asian house shrew (Suncus murinus) was remarkably abundant
in primary forest at high elevation; this species has also been found to be abundant in
montane primary forest on Negros Island, which also has low total species richness. Species
richness of small nonflying mammals was greatest at fairly high elevation.

1 Field Museum of Natural History, 1400 South Introduction
Lake Shore Drive. Chicago. IL 60605-2496. U.S.A.

2 Department of Biology, Iligan Institute of Tech- Tne Philippine Islands present a remarkable
nology, Mindanao State University, Iligan City, theater for the study of the ecology and evolution
Lanao del Norte. Philippines. of mammalian diversity. Its islands range in size

1 Laksambuhay Conservation. Inc., 10241 Mt. from less than one hectare to over 100,000 km2,

Bulusan Street, U-2. Los Banos, Laguna. Philippines. with geological age varying from under 1 million

28 FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, PP. 28-48

years to over 40 million years. These islands
represent many sets of historically distinct geo-
logical units of remarkably varied origins; some
had land-bridge connections to the Asian main-
land in the past (those of the Palawan group),
but most are purely oceanic in origin (Heaney,
1986, 1991b, 2000; Heaney & Rickart, 1990;
Hall, 1998, 2002). The mammals that have
evolved in this diverse archipelago include at
least 1 1 1 species endemic to the archipelago out
of 172 native terrestrial species; with endemism
at 64%, the Philippine fauna is one of the most
distinctive in the world (Mittermeier et al., 1997,
1999; Heaney & Regalado, 1998). While most of
the endemic species occur on the large islands of
Luzon, Mindanao, Mindoro, Negros, and Pala-
wan (e.g., Heaney, 1986, 1993, 2000; Heaney et
al., 1998; Rickart et al., 1998), significant
numbers occur on the smaller islands as well,
especially those surrounded by deep water (e.g.,
Heaney, 1986, 2004; Goodman & Ingle, 1993;
Heaney & Tabaranza, 1997; Musser et al., 1998).
As noted by Heaney and Tabaranza (2006a),
Camiguin, an island of 265 km2 located about
10 km north of Mindanao in the Bohol Sea, is
one such deep-water island, with a minimum
depth to Mindanao of 385 m. It is steeply
mountainous, with several active volcanic cones
that reach to a maximum elevation of about
1600 m. A series of biological surveys on
Camiguin in the late 1960s that focused on birds
(see Balete et al., 2006) also yielded some
mammal specimens, and an earlier report on
those surveys (Heaney, 1984) concluded that the
island had no endemic mammal species and was
depauperate. Subsequent studies on other islands
made us suspect that those earlier surveys were
incomplete because so few mammal species had
been obtained and because the number of
nonvolant mammal specimens was small (thus
indicating limited sampling effort). Further, on
the basis of biogeographic patterns elsewhere in
the Philippines, we predicted the presence of
several endemic small mammals on Camiguin
(Heaney, 2004). To investigate the hypotheses
that the previously measured species richness was
low because of incomplete surveys and that
about two endemic species should be present, we
returned to Camiguin briefly in 1992 and more
extensively in 1994 and 1995 to conduct addi-
tional mammal inventories in all the major
habitats along the elevational gradient, especially
by trapping small mammals at higher elevations
where there were few records from the 1 960s. As

indicated in a brief preliminary report (Heaney &
Tabaranza, 1997), we found eight additional
species on the island that are widespread in the
Philippines, plus two previously unknown en-
demic species of rodents. The purpose of this
paper is to summarize the results of the 1994 and
1995 mammal surveys and integrate those data
with information from the 1960s, including
information on habitat associations, relative
abundance, and ecology of the species.

Methods
Prior Reports

The first report of mammals from Camiguin
Island was that of Gray (1843), who reported
Paradoxurus hermaphroditus. Three field teams
from Silliman University led by Dioscoro S.
Rabor collected mammals on Camiguin in 1967,
1968, and 1969; specimens were deposited at the
Delaware Museum of Natural History (dmnh)
and Royal Ontario Museum (rom); for details,
see Heaney and Tabaranza (2006a). Several
specimens were reported by Peterson and Fenton
(1970); all known specimens from the 1960s were
examined and summarized by Heaney (1984). All
data included in this paper from the 1960s
specimens are based on data in Heaney (1984),
except as noted below.

Recent Data

Field studies were conducted during three
periods in 1992, 1994, and 1995; general methods
and site descriptions are given in Heaney and
Tabaranza (2006a). Sampling during 1995 fol-
lowed methods used on Leyte, Luzon, Negros,
and other islands (Heaney et al., 1989, 1999;
Rickart et al., 1991, 1993) to facilitate quantita-
tive comparisons. Nonvolant small mammals
were caught in traps; during 1995, all traps were
Victor rat snap traps. Most were baited with
fresh fried coconut coated with peanut butter,
but a few were baited with live earthworms.
During 1994, several National live traps were
used, in addition to Victor rat traps, and were
baited with coconut bait. Bats were captured in
12-m mist nets. Voucher specimens were pre-
pared in fluid or as skeletons and have been
deposited at The Field Museum of Natural
History (fmnh), National Museum of the Phi-

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

29

lippines (nmp), and Mindanao State University-
lligan Institute ol' Technology (msu-iit). Most
specimens were autopsied for reproductive in-
formation. The si/e of embryos was measured as
crown to rump length (CRL). Subadult animals
are defined here as those that have not completed
cranial growth, especially those having unfused
basicranial sutures; these young animals have
pelage that is usually softer and grayer than that
of adults and are noticeably lower in weight and
females are usually nulliparous. Young adults
are older; they have nearly completed cranial
growth but have not yet reached adult weight
and, usually, have not yet reproduced or are
pregnant for the first time. Adults have complet-
ed cranial growth and adult pelage, and usually
the females are multiparous. Comments on
distribution and use of scientific names are based
on Heaney et al. (1998) unless additional sources
are mentioned. Records of specimens examined
are summarized at the end of each account; such
summaries include site number and the number
of specimens (in parentheses).

External measurements and weights reported
here were taken in the field by members of
the field team on fresh animals. Cranial mea-
surements were taken by Heaney with digital
calipers graduated to 0.01 mm. Comparisons of
cranial measurements are to published records of
specimens measured in the same manner by
Heaney.

Accounts of Species

Order Insectivora

Family Soricidae — Shrews

Crocidura beatus Miller, 1910

The Mindanao shrew is widespread on islands
in the Mindanao Faunal Region (Heaney &
Ruedi, 1994; Heaney et al., 1998); these are the
first records from an island that was not part of
the late Pleistocene island of Greater Mindanao
(Heaney. 1986). This shrew has most often been
found in primary forest, especially at higher
elevations; is usually uncommon in secondary
forest; and is absent outside of forest (Heaney et
al.. 1989; Rickart et al., 1993).

Crocidura beatus was trapped on Camiguin at
three forest sites in May 1994 and March 1995
(Fig. 1. Table 1). It was uncommon in secondary
lowland forest at 1000 m elevation (Site 4), in
disturbed lower montane forest at 1200 m

elevation (Site 5), and in primary montane forest
at 1275 m (Site 7). It was most often trapped
under tree roots and live vegetation. None were
taken in agricultural areas at 150 m (Site 3), or in
mossy forest at 1475 m (Site 8, where there was
limited sampling; Table 1 ). This use of habitat is
consistent with data from islands on Greater
Mindanao (Rickart et al., 1993; Heaney et al.,
unpubl. data).

In 1994 and 1995, three adult females were
trapped; one was pregnant with a single embryo
(CRL = 10 mm). A multiparous, nonpregnant
female weighed 13 g; an adult male weighed 7 g.
Both cranial and external measurements
(Table 2) are within stated ranges for Mindanao
(Heaney & Ruedi, 1994) but are slightly smaller
than those of series taken on Biliran, Leyte, and
Maripipi (Rickart et al., 1993).

Specimens Examined — Total 5. Site 4 (2 fmnh,
1 msu-iit); Site 5 (1 fmnh); Site 7 (1 fmnh).

Suncus murium (Linnaeus, 1 766)

The Asian house-shrew occurs widely in Asia
and Indo-Australia; it now occurs throughout
the Philippines, though it is not native to the
country. It is abundant in urban and agricultural
areas; on islands with low mammal species
richness such as Negros, it is sometimes abun-
dant in disturbed forest and occasionally in
primary forest (Heideman et al., 1987; Heaney et
al., 1989, 1991), but on islands of average species
richness, it is usually rare or absent from forest
(Heaney et al., 1989, 1998. unpubl. data; Rickart
et al., 1993).

A single subadult specimen from Mt. Tim-
poong Peak was available previously (Heaney
1984). In 1994-1995, we captured this species
from 150 to 1475 m, and it was the most
common species at the three highest sites, all in
primary forest (Table 1, Fig. 1). It was especially
abundant in montane forest at 1275 m (Site 7). It
was moderately abundant in primary mossy
forest (Site 6; elev. 1300 m) and in lower mossy
forest at 1475 m elevation (Site 8) but was much
less common in heavily disturbed lowland
agricultural land at 150 m (Site 3). This pattern
of abundance is quite different from that on the
species-rich islands of Biliran, Leyte, Luzon,
Maripipi, and Mindanao, where specimens were
never caught in primary forest (Heaney et al.,
1989, 1999, unpubl. data; Rickart et al., 1993)
but similar to the species-poor island of Negros,
where S. murinus was abundant in transitional
mossy/montane forest and in mossy forest at

30

FIELDIANA: ZOOLOGY

1800 i-
1600 ="
1400 —

~ 1200

O 1000 —

re _

| 800

600 —
400 —
200

0

*=1960s
■=1990s

11

•a 5)

o

maximum elevation

J 1 1 1. ■ 1 1

L "

o .c
c £

t c

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II

5 c
o cc

II

0)

</> c/>
5 ^

mossy

montane

lowland

primary
disturbed

no forest

II

c

"3

Fig. 1. Elevational range of nonvolant small mammals (Insectivora and Rodentia) on Camiguin Island,
Philippines. Records from the 1960s are indicated with stars and from the 1990s by solid squares. The approximate
original boundaries of primary lowland, montane, and mossy rain forest along the elevational gradient are
indicated. The condition of forest along our transect in the middle 1990s is indicated as nearly absent (below
600 m), disturbed by logging and agriculture but present as second growth (about 600-1250 m), and primary or
lightly disturbed by human activities and landslides (above about 1250 m). Elevations from the 1960s were rough
estimates (see text).

1280 m (Heaney et al., 1989), which is similar to
Sites 6 and 7.

Suncus murinus was most often trapped in
runways or clear areas beneath fallen and rotting
logs, under roots of trees, or under horizontal
trunks of live trees as well as in runways near
large boulders. Many were caught during day-
light hours.

Five adult females with a mean weight of 32 ±
4.5 g (range = 27-39 g) were pregnant; litter
sizes for four of these were one, two, three, and
three. Thirteen nonpregnant parous females
(those with large mammae) weighed an average
of 28.8 ± 4.1 g (range = 22-35 g), and nullipa-
rous females (those with small mammae)
weighed 20.6 ± 2.2 g (range = 17.5-23 g, N =
10). Adult males (defined as those with large
testes) had a mean weight of 36 ± 5.8 g (range
24-48 g, N = 27). Males are conspicuously
larger than females in this species (Table 2).

Specimens Examined — Total 78. Site 3 (2 msu-
iit); Site 6(10 msu-iit); Site 7 (56 fmnh); Site 8 (9
fmnh); Site 15 (1 dmnh).

Order Chiroptera

Family Pteropodidae — Fruit Bats

Cynopterus brachyotis (Muller, 1838)

The common short-nosed fruit bat is wide-
spread in Southeast Asia and is common
throughout the Philippines. It ranges from sea
level to at least 1250 m and is typically found in
agricultural areas; it is also common in second-
ary lowland forest but usually rare in primary
forest (Heaney et al., 1998).

Our limited netting on Camiguin during 1994-
1995 (Table 3) indicated that C. brachyotis was
abundant in a highly disturbed lowland agricul-
tural area at 10 m elevation (Site 1) and was
common in a heavily disturbed lowland agricul-
tural area at 100 m (Site 2; Table 3). It was less

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

31

Tabu 1. Numbers of nonvolant small mammals captured in traps in heavily disturbed lowland agricultural
area (Site 3) secondary lowland forest (Site 4). disturbed lower montane forest (Site 5), primary mossy forest (Site
6) primary montane forest (Site 7), and lower mossy forest (Site 8) on Camiguin Island. The numbers of captures
per KM) trap-nights are given in parentheses. See Heaney and Tabaranza (2005a) for full site descriptions. Asterisks
mark species presumed to be present; see Methods.

Site 3,

Site 4,

Site 5,

Site 6,

Site 7,

Site 8,

Scientific name

150 m

1000 m

1200 m

1300 m

1275 m

1475 m

Total

Crocidura bcutits

0

3 (0.3)

1 (0.3)

0*

1 (0.2)

0

5

Sanctis marinas

2(1.4)

0*

0*

10(2.9)

56(8.5)

9 (2.3)

77

Apomys camiguinensis

0

14(1.5)

0*

2 (0.6)

9(1.4)

0

25

Bullimus gatnay

0

4 (0.4)

2 (0.6)

3 (0.9)

10(1.5)

1 (0.3)

20

C 'runomys melanius

0

2 (0.2)

1 (0.3)

0*

2 (0.3)

0

5

Rattas everetti

0

7(0.8)

2 (0.6)

3 (0.9)

2 (0.3)

2 (0.5)

16

Rat t us exulans

2(1.4)

0*

0*

0*

5 (0.8)

0

7

Rattas tanczumi

21 (14.2)

8 (0.9)

4(1.2)

1 (0.3)

0

0

34

Total small mammals

24

38

10

19

85

12

188

Total trap-nights

148

907

339

348

655

386

2783

Number/ 100 trap-nights

16.2

4.2

2.9

5.5

9.9

3.1

6.8

Total small mammal species

3

6(+l)

5 (+3)

5 (+3)

7

3

8

Native small mammal species

0

5

4(+l)

3 (+2)

5

2

5

common in disturbed lowland forest at 1000 m
(Site 4) and uncommon in disturbed lower
montane forest at 1000-1300 m and mossy forest
between 1200 and 1400 m elevation (Sites 5 and
6). Limited netting did not detect this species in
montane primary forest at 1275 m elevation (Site
7). The occurrence of C. brachyotis in these
habitats is consistent with records from other
Philippine islands; for example, on Catanduanes,
Leyte, Luzon, and Negros, this species was
most abundant in agricultural land and second-
ary forest (Heaney et al., 1989, 1991, 1999;
Heideman & Heaney, 1989; Ingle, 1992; Rickart
et al., 1993), similar to Sites 1 and 3 to 5 on
Camiguin. Records from the 1960s indicate that
the species occurs along the entire elevational
gradient, from sea level to near the peaks,
although most specimens are from below
1000 m (i.e., below the transition to montane
forest; Fig. 2).

Eight adult females (mean = 29.5 ± 4.72 g)
taken in May 1992 and 1994 were pregnant with
single embryos (CRL = 5-26 mm). Two of the
pregnant females (22 and 35 g) and three non-
pregnant ones (32.5-39 g) were lactating. Adult
females with enlarged mammae but not pregnant
or lactating had mean weight of 29.2 ± 2.3 g
(range = 26 33.2 g, N = 13). Five nulliparous
females had a mean weight of 16.5 g ± 2.0 g
(range = 14-19 g). Males with abdominal testes
had a mean weight of 27.4 ± 2.6 g (range = 22-
32.5 g, N = 15). On Negros Island, this species
probably produces two young per year, one in

March/April and another in August/September
(Heideman, 1995).

Females are slightly larger than males in most
external and cranial dimensions (Table 4), a trend
quite similar to specimens from Mt. Kitanglad,
Mindanao (Heaney et al., unpubl. data). Speci-
mens from Biliran, Leyte and Maripipi, in
contrast, showed the opposite trend (Rickart et
al., 1993).

Specimens Examined — Total 91. Site 1 (27
fmnh); Site 3 (5 msu-iit); Site 4 (4 fmnh, 14 msu-
iit); Site 6 (1 fmnh, 3 msu-iit); Site 11(12 dmnh);
Site 12 (3 dmnh); Site 13 (6 dmnh); Site 16 (2
rom); Site 17 (14 rom).

Harpyionycteris whiteheadi Thomas, 1 896

The harpy fruit bat has been reported pre-
viously from Camiguin (Peterson & Fenton,
1970), as well as from Marinduque, Masbate,
Mindoro, Negros, southern Luzon, and through-
out Greater Mindanao (Heaney et al., 1998,
1999). It is generally restricted to primary or
lightly disturbed forest; it is usually rare in
lowland forest but often is moderately common
in montane forest from roughly 800 m to at
least 1800 m. It apparently feeds heavily on
the fruits of viney pandans (Freycinetia spp.) and
figs (Ficus spp.); (Heaney et al., 1989, 1999;
Heideman & Heaney, 1989; Rickart et al.,
1993).

In May 1992, we netted one pregnant female
weighing 123 g with a single embryo (CRL =
28 mm) at 10 m elevation, in a lowland

32

FIELDIANA: ZOOLOGY

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agricultural area (Site 1; Table 3). On Mind-
anao (Kitanglad Range), pregnant females
were similarly recorded in May, with further
records of pregnancy in April, August, and
October; lactation was noted in March, April,
and August (Heaney et al., unpubl. data). In
March 1995, at Site 7 (1275 m in primary
montane forest), where Freycinetia spp. were
abundant, we often heard the distinctive Har-
pyionycteris whistles (Rickart et al., 1993; Hea-
ney et al., 1999), but none were captured.
Combined with records from the 1960s (Fig. 2),
these data indicate that it occurs on this island
from sea level to 1500 m in lowland, montane,
and mossy forest.

External and cranial measurements of H.
whiteheads on Camiguin are comparable to those
from Mindanao (Mt. Kitanglad) but are slightly
larger than those found on Leyte and Luzon
(Mt. Isarog) (Heaney, 1984; Rickart et al., 1993;
Heaney et al., 1999, unpubl. data).

Specimens Examined — Total 4. Site 1 (1
fmnh); Site 11 (1 dmnh); Site 12 (1 dmnh); Site
16 (1 ROM).

Macroglossus minimus (E. Geoffroy, 1810)

The dagger-toothed flower bat occurs from
Thailand to Australia and is found throughout
the Philippines (Heaney et al., 1998). Within the
Philippines, it occurs in virtually every habitat in
the country, from sea level to at least 2250 m. It
is often abundant in agricultural and heavily
disturbed areas, is common in secondary forest,
and usually is uncommon in primary forest. It is
most often associated with domestic or wild
banana (Musa spp.; Heaney et al., 1989, 1999;
Heideman & Heaney, 1989; Rickart et al.,
1993).

On Camiguin, our limited netting showed M.
minimus to be abundant in a highly disturbed
lowland agricultural area at ca. 10 m elevation
(Site 1), present in a heavily disturbed lowland
agricultural area at 100 m (Site 3), uncommon in
disturbed lowland forest at 1000 m (Site 4),
common in disturbed lower montane forest at
1000-1300 m (Site 5), and present in mossy
forest between 1200 and 1400 m elevation (Site
6; Table 3). Limited netting did not detect this
species in primary montane forest at 1275 m
elevation (Site 7). Combined with prior records,
it is apparent that this species occurs on
Camiguin from sea level to at least 1200 m, in
lowland and montane forest, in both disturbed
and undisturbed forest (Fig. 2).

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

33

Tabu: 3. Numbers of fruit bats captured in mist nets in a lowland agricultural area (Site 1), heavily disturbed
lowland agricultural area (Site 3), secondary lowland forest (Site 4), disturbed lower montane forest and primary
mossy forest (Sites 5 and 6 concurrently), and primary montane forest (Site 7) on Camiguin Island during 1992,
1994, and 1995. The number of captures per net-night are given in parentheses. See Heaney and Tabaranza (2006a)
for full site descriptions. Asterisks indicated species observed but not netted (see text).

Scientific name

Site I,
10 m

Site 3,
150 m

Site 4,
1000 in

Sites 5 and 6,
1200-1400 m

Site 7,
1275 m

( 'ynopterus brach vol is

27 (4.5)

5(1.67)

18(0.75)

4 (0.29)

0

liar prion vcteris whitehead!

1 (0.17)

0

0

0

0*

Macroglossus minimus

15 (2.5)

1 (0.33)

4(0.17)

9 (0.64)

0

Picnot hints jagori

8(1.33)

0

16(0.67)

1 (0.07)

0

Total fruit bats

51

5

35

14

0

Total net-nights

6

3

24

14

8

Fruit bats per net-night

8.5

1.67

1.46

1.00

0

Six adult females netted in May 1992 and
1994. with a mean weight of 15.2 ± 3.2 g (range
= 12.5-20.3 g), were pregnant with single
embryos. Four parous females with large mam-
mae but neither pregnant nor lactating had
a mean weight of 18.5 ± 1.78 g (range = 16-
20 g). Three adult males weighed between 16 and
19 g. A juvenile male and a juvenile female each
weighed 5 g. On Mindanao (Kitanglad Range),
pregnant females were recorded in April to

August and October, while lactating females
were recorded in May, September, and October
(Heaney et al., unpubl. data). Heideman (1995)
documented that this species undergoes aseaso-
nal breeding and postpartum estrus on Negros
Island and has several young per year.

Comparison with specimens of M. minimus
from Biliran, Dinagat, Leyte, Luzon (Mt.
Isarog), Maripipi, and Mindanao (Kitanglad
Range) shows that while the overall variation

1800

1600

1400

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§ 1000

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600

400
200

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■=1990s

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indicates a documented record without a voucher specimen (see text).

34

FIELDIANA: ZOOLOGY

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HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

35

in cranial and external measurements is slight,
Camiguin specimens tend to cluster consistently
within the upper ranges of most features
measured (Table 4; Heaney and Rabor, 1982;
Heaney et al., 1991, 1999, unpubl. data; Rickart
et al., 1993).

Specimens Examined — Total 56. Site 1 (15
fmnh); Site 3 ( 1 msu-iit); Site 4 (4 msu-iit); Site 5
(8 msu-iit); Site 6 (1 msu-iit); Site 11(12 dmnh);
Site 13 (5 dmnii); Site 14 (2 dmnii); Site 17 (8
dmnii).

Ptenochirus jagori ( Peters, 1861)

The musky fruit bat is a common Philippine
endemic, occurring throughout the archipelago
with the exception of the Batanes/Babuyan and
Palawan faunal regions from sea level to at least
1800 m (Heaney et al., 1998). Our limited netting
on Camiguin showed Ptenochirus jagori to be
common in a lowland agricultural area at 10 m
elevation (Site 1), less common in disturbed
lowland forest at 1000 m (Site 4), and scarce in
primary mossy forest at 1300 m (Site 6; Table 3).
Combined with records from the 1960s, these
data indicate that the species is widespread in
lowland and montane rain forest from sea level
to at least 1200 m (Fig. 2), though probably its
abundance declines with increasing elevation and
with increasing levels of disturbance (Table 3), as
noted elsewhere (Heaney et al., 1989, 1999,
unpubl. data; Heideman & Heaney, 1989; Rick-
art et al., 1993; Lepiten, 1997).

Three adult females, netted in May 1992 and
1994, weighing an average of 70 g (range = 68-
75 g), were pregnant with a single embryo each
(CRL = 5 10 mm). Three nonpregnant females
with large mammae had an average weight of
73.5 g (range = 72-74 g), and two nulliparous
females weighed 67 and 68.5 g. Eleven adult
males had a mean weight of 73.2 ± 4.7 g (range
= 64-78 g, N = 11). Pregnant females of P.
jagori have been recorded also in May on Luzon
(Mt. Isarog) (Heaney et al., 1999). On Mindanao
(Kitanglad Range), pregnant females were re-
corded in March, May, July, and August and
lactating females in May to June and August to
December (Heaney et al.. unpubl. data). Heide-
man and Powell (1998) found that on Negros
Island, P. jagori gives birth to a single young
twice each year: the first in late March or early
April and the second in August. It was further
discovered that this species undergoes delayed
implantation and early development that lasts
for five months, shorter than in two other

endemic species of Philippine pygmy fruit bats,
Haplonycteris fischeri and Otopteropus eartilago-
nodus, where the phenomenon was first detected
(Heideman, 1989; Heideman et al., 1993; Heide-
man & Powell, 1998). Additionally, this condi-
tion was apparently exhibited by primiparous
young adult females only, allowing them to give
birth only once in their first year, which had the
effect of enabling them to synchronize breeding
with the adult females the following year (Heide-
man & Powell, 1998).

Males are somewhat larger than females in
most cranial and external dimensions, as on
Biliran, Leyte, and Maripipi (Table 4; Rickart et
al., 1993). Cranial and external measurements
(Table 4) are noticeably larger than those for
series from Catanduanes and southern Luzon
and were similar to those from Biliran, Leyte,
and Maripipi (Heaney, 1984; Heaney et al., 1991,
1999; Rickart et al., 1993).

Specimens Examined — Total 46. Site 1 (8
fmnh); Site 4(16 msu-iit); Site 6 (1 msu-iit); Site
11 (6 dmnh); Site 13 (6 dmnh); Site 17 (9 rom).

Pteropus hypomelanus Mearns, 1905

The common island flying fox occurs from
Thailand to Australia and is found throughout
the Philippines with the exception of the Palawan
faunal region. It is often common in agricultural
areas from sea level to ca. 900 m and is absent in
primary forest (Heideman & Heaney, 1989;
Heaney et al., 1991, 1998; Utzurrum, 1992;
Rickart et al., 1993). Records from the 1960s
document it from Camiguin at elevations from
about 250 to 1500 m (Fig. 2), but we netted none
in the 1990s; because this species typically flies
above the canopy and our nets were set not more
than about 4 m above the ground, our failure to
catch any does not necessarily indicate any
change in their abundance.

External and cranial measurements show only
slight variations with those of specimens from
Dinagat, Panay, Leyte, and Maripipi (Heaney &
Rabor, 1982; Rickart et al., 1993).

Specimens Examined — Total 8. Site 10 (2
dmnh); Site 12 (1 dmnh); Site 13 (2 dmnh); Site

17 (3 ROM).

Pteropus pwnilus Miller, 1910

The little golden-mantled flying fox is endemic
to the Philippines (aside from a single population
on Miangas Island. Indonesia, adjacent to
Mindanao) and occurs throughout the archipel-
ago, with the exception of the Batanes/Babuyan

36

FIELDIANA: ZOOLOGY

and Palawan regions (Heaney et al., 1998).
Previously reported from Camiguin as P. tablasi,
the species was revised to include P. tablasi and
P. balutus as synonyms under Pteropus pumilus
(Klingener & Creighton, 1984). It is associated
with primary and good secondary lowland forest
from sea level to about 1100 m, and it is
uncommon outside of forest. Additionally, it is
most common on small islands and is uncommon
to rare on larger islands. Pteropus pumilus often
is netted in clearings or on ridgetops (Heideman
& Heaney, 1989; Utzurrum, 1992; Rickart et al.,
1993).

We did not encounter P. pumilus during the
1990s, but the 1967-1969 surveys obtained 42
individuals at four sites from ca. 250 to nearly
1000 m elevation (Fig. 2). As with Pteropus
hypomelanus, this species usually flies above the
canopy, so our failure to catch any in the 1990s
does not necessarily indicate a change in their
status on the island.

Specimens Examined — Total 42. Site 11 (19
dmnh); Site 13 (2 dmnh); Site 14 (1 dmnh); Site

17 (20 ROM).

Family Emballonuridae — Sheath-tailed Bats

Emballanura alecto (Eydoux and Gervais, 1836)
The Philippine sheath-tailed bat is a common
cave-dwelling species that occurs throughout
most of the Philippines and is also known from
Borneo, the Moluccas, and Sulawesi (Koopman,
1989). It has been recorded only in lowland areas
(450 m and below) in disturbed forest and
agricultural areas with scattered remnant forest,
with most captures recorded from caves, under
large boulders, or in man-made tunnels (Heaney
et al., 1991, 1999; Rickart et al., 1993). We did
not record this species during our fieldwork in
1994 and 1995, but in 1967 specimens were taken
from Tag-ibo Cave at 400 ft (ca. 120 m) and at
1400-3300 ft (ca. 400-1000 m) on Mt. Mamba-
jao (see also Heaney, 1984).

Comparison of external and cranial measure-
ments with series from Leyte and Biliran shows
little variation (Table 4; Rickart et al., 1993).

Specimens Examined — Total 4. Site 17 (2
rom); Site 18 (2 rom).

Family Rhinolophidae — Horseshoe-nosed Bats

Rhinolophus arcuatus Peters, 1871

The arcuate horseshoe bat is widespread from
Sumatra to New Guinea and throughout the
Philippines (Heaney et al., 1998), including the
Palawan faunal region (Esselstyn et al., 2004). It

is most often encountered from lowlands to at
least 1350 m in agricultural lands to primary
lowland and montane forest and occasionally
roosts in caves (Heaney et al., 1991, 1999;
Rickart et al., 1993). Two groups that differ in
size and habitat are recognized within this
species: a smaller morphotype, designated R.
arcuatus — s, that occurs in the lowlands or
disturbed habitats, and a larger one, designated
R. arcuatus — 1, found in forest at higher eleva-
tions (Ingle & Heaney, 1992).

In May 1994, we netted this species at 1000 m
elevation in disturbed lowland forest (Site 4). Of
two adult females netted, one (15 g) was
pregnant with one large embryo (CRL =
28 mm), and the other was lactating (12.5 g).
Pregnancies in this species were recorded also in
March on Luzon (Mt. Isarog) and in April on
Biliran, Leyte, and Maripipi (Rickart et al., 1993;
Heaney et al., 1999).

Cranial and external measurements (Table 5)
of the Camiguin specimens are consistently
larger than those in series from Biliran, Leyte,
Luzon (Mt. Makiling), Maripipi, and Mindanao
(Mt. Kitanglad), which all fall within the
dimensions of the smaller morphotype, R.
arcuatus — s (Ingle & Heaney, 1992; Rickart et
al., 1993; Heaney et al., 1999, unpubl. data).
Instead, their external and cranial dimensions are
comparable to the larger morphotype, R. arcua-
tus— 1 (Ingle & Heaney, 1992). Systematics in this
"species" are badly in need of detailed study.

Specimens Examined — Total 10. Site 4 (4 msu-
iit); Site 13 (6 dmnh).

Rhinolophus inops K. Anderson, 1905

The Philippine forest horseshoe bat is common
to abundant in primary lowland and montane
forest from sea level to 2250 m and is usually
rare in secondary forest and mossy forest
(Heaney et al., 1991, 1998, 1999; Rickart et al.,
1993). Improvements in our understanding of
Philippine Rhinolophus lead us to reidentify the
single specimen from Site 13, reported as R.
subrufus by Heaney (1984), as R. inops. We
netted two additional males in disturbed lowland
forest at 1000 m (Site 4). Cranial and external
measurements (Table 5) are slightly larger in
most dimensions than those of the series from
Biliran, Leyte (Rickart et al., 1993), and Mind-
anao (Mt. Kitanglad; Heaney et al., unpubl.
data); we suspect that regional morphs corre-
sponding to the Pleistocene islands (Heaney,
1986) are present.

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

37

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Specimens Examined — Total 3. Site 4 (2 msu-
lii); Site 13 (1 dmnh).

Rhinolophus virgo K. Anderson, 1905

The yellow-faced horseshoe bat is an endemic
species widespread within the Philippines. It is
usually encountered in primary lowland
forest from 250 to 1100 m; several records are
from caves (Heaney et al, 1991; Rickart et al.,
1993). We obtained one adult female from
the edge of a small tree-fall gap in primary
montane forest at 1275 m on 22 March 1995
(Site 7). It has small mammae and showed no
indication of reproductive activity. Cranial
dimensions (Table 5) fall within the range of
those from Leyte and Maripipi but are slightly
smaller than those from Mindanao (Mt. Kitan-
glad; Rickart et al, 1993; Heaney et al., unpubl.
data).

Specimens Examined — Total 1. Site 7 (1
fmnh).

Family Vespertilionidae — Common Bats

Murina cyclotis Dobson, 1872

The round-eared tube-nosed bat is widespread
in southern Asia and was previously known in
the Philippines from a few specimens from the
central and southern portion of Greater Luzon
as well as Greater Mindanao, Sibuyan, and
Siquijor, with records from disturbed and
primary lowland and montane forest from 250
to 1500 m (Heaney et al., 1991, 1998, 1999,
unpubl. data; Rickart et al., 1993; Ruedas, 1994;
Lepiten, 1997). We netted one adult female in
May 1994 in secondary lowland forest at 1000 m
(Site 4).

Specimens Examined — Total 1. Site 4 (1 msu-
iit).

Pipistrellus javanicus (Gray, 1838)

The Javan pipistrelle is found from Korea
to Java and throughout the Philippines. It is
generally common in primary montane forest
and uncommon in primary lowland and mossy
forest, though it has been found from sea
level to 2250 m (Ingle, 1992; Heaney et al.,
1998, 1999, unpubl. data). A single skin without
a skull, tentatively identified as this species, was
obtained from Camiguin by the 1960s surveys. In
March 1995, two were netted in primary
montane forest at 1275 m (Site 7). Cranial and
external measurements (Table 5) are comparable
with those of a series from Mindanao (Mt.
Kitanglad) in most dimensions (Heaney et al.,
unpubl. data).

38

FIELDIANA: ZOOLOGY

Specimens Examined — Total 3. Site 7 (2
fmnh); Site 13 (1 dmnh).

Order Primates

Family Cercopithecidae — Monkeys

Macaca fascicularis (Raffles, 1821)

The long-tailed macaque occurs from Burma
to Timor and is found throughout the Philip-
pines (Fooden, 1995; Heaney et al., 1998). It is
found in agricultural areas near forest, in second-
growth and secondary forest, as well as primary
forest from sea level to at least 1800 m in
lowland and montane forest (Goodman & Ingle,
1993; Rickart et al., 1993; Heaney et al., 1999).
We did not collect any specimens, but in March
1995 we obtained the skull of an adult individual
from a hunter who had killed the macaque in
a forested area above Mahinog a few years
earlier. Also in March 1995, a local hunter who
was a member of our team saw two macaques
a short distance from our camp at 1100m
elevation (Site 7) in primary montane forest.
He reported that the macaques were fairly
common in what remained of lowland primary
forest and good secondary forest at that time and
were actively hunted.

Specimens Examined — Total 1. Above Mahi-
nog (1 fmnh).

Order Rodentia

Family Muridae — Rats and Mice

Apomys camiguinensis Heaney and Tabaranza,
2006

A previously unknown species of Apomys
(Musser, 1982) was trapped on Camiguin in
May 1994 and March 1995 (Heaney & Tabar-
anza, 1997) and described in this volume
(Heaney & Tabaranza, 2006b). The Camiguin
forest mouse is moderately large for the genus,
averaging about 40 g. It is marked by a number
of subtle morphological characters, and molec-
ular data show it to be most closely related to
Apomys hylocoetes and A. insignis from Mind-
anao and to an unnamed species from Leyte and
Biliran (Steppan et al., 2003; Heaney & Tabar-
anza, 2005b). This species was common (Table 1,
Fig. 1) in disturbed lowland forest at 1000 m
(Site 4) and in primary montane forest at 1275 m
(Site 7) and relatively uncommon in primary
mossy forest at 1200-1400 m (Site 6). It was
not found in heavily disturbed lowland agri-
cultural land (Site 3, elev. 150 m), lower
montane forest (Site 4, elev. 1000-1300 m), or
lower mossy forest (Site 8, elev. 1475 m), but

there was limited sampling at all of these
(Table 1). Most specimens were taken from traps
placed in primary montane forest underneath
root tangles and beneath fallen and rotten logs
on a moderate slope. All individuals were
captured at night.

A pregnant female taken on 29 May 1994
weighed 40.5 g and had a single embryo (CRL =
30 mm). Adult females with large mammae but
not pregnant weighed 39 ± 2.1 g (range = 37.5-
42 g, N = 5); two of these, taken 23 May 1994
and 17 March 1995, each had two placental
scars. Nulliparous females (with small mammae)
were lighter and had an average weight of 35 g
(range = 34-36). Females have two pair of
inguinal mammae. These data indicate a litter
size between one and two. Males with scrotal
testes had a mean weight of 41.3 ± 4.7 g (range
= 36.5^48 g, N = 11); testis size ranged from 15
X 16 mm to 8 X 12 mm. Males with abdominal
testes weighed 33.0 ± 5.0 g (range = 28-34 g, N
= 5) and had testis size of 5 X 8 mm.

Specimens Examined — Total 25. Site 4 (14
msu-iit), Site 6 (2 msu-iit); Site 7 (9 fmnh).

Bullimus gamay Rickart, Heaney and Tabaranza,
2002

A previously unknown species of Bullimus,
recently described as B. gamay (Rickart et al.,
2002), was captured on Camiguin in May 1994
and March 1995. The Camiguin forest rat was
incorrectly assigned to Tarsomys by Heaney and
Tabaranza (1997) and was designated "Bullimus
sp. A" by Heaney et al. (1998). It was trapped
from 900 to 1475 m (Fig. 1, Table 1) but not in
heavily disturbed lowland agricultural land from
sea level to 300 m (Site 3). Most specimens were
taken from traps placed in runways beneath root
tangles and rotten logs or near large rocks. This
species was most common in primary montane
forest at 1275 m (Site 7). It was less common in
disturbed lower montane forest at 1000-1300 m
(Site 5) and in primary mossy forest at 1200-
1400 m (Site 6). It was rare in secondary lowland
forest at 1000 m (Site 4) and in lower mossy
forest at 1475 m (Site 8). Bullimus bagobus
occurs in similar habitat on Mindanao but also
occurs at lower elevation where good forest
persists (Heaney, 2001; Heaney et al., unpubl.
data); this suggest that B. gamay should also be
sought at lower elevations on Camiguin.

Parous females with large mammae weighed
an average of 370 g (range = 360-390 g, N = 3),
while nulliparous young adult females with small

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

39

mammae were lighter, having a mean weight of
276 ± 53.7 g (range = 210-345 g, N = 5). Adult
males with scrotal testes had an average weight
of 402 g (range = 240-500 g; N - 3) and had
testis size ranging from 21 x 30 mm to 27 X
35 mm). Among the nine individuals taken in
May 1994, the smallest was a nulliparous young
female weighing 210 g, while among the 11
individuals captured in March 1995, two were
juveniles, a male weighing 120 g and a female
weighing 125 g. No pregnant females were taken
in May 1994 and March 1995, but a lactating
female (360 g) was taken on 18 March 1995, and
a female taken on 29 May 1994 had three
placental scars. This suggests that females give
birth in February and March. Three females had
three pairs of mammae and one had four pairs,
indicating that litter size most likely does not
exceed three.

Specimens Examined — Total 20. Site 4 (4 msu-
iit); Site 5 (2 msu-iit); Site 6 (3 msu-iit); Site 7 (10
emnh); Site 8 (1 fmnh).

Crunomys melanius Thomas, 1907

The Mindanao shrew-mouse was previously
known only from Leyte and Mindanao from
near sea level to 1550 m, apparently in primary
lowland rain forest; it is rare in collections
(Musser & Heaney, 1992; Heaney et al., 1998;
unpubl. data; Rickart et al., 1998). On Camiguin,
we found it to be uncommon in heavily disturbed
lowland agricultural land at 1000 m (Site 4), in
disturbed lower montane forest at 1200 m (Site
5), and in primary montane forest at 1275 m
(Site 7; Table 1, Fig. 1). It was trapped beneath
rotten logs and wood tangles in an area with few
dead leaves and little moss. It occurs down to sea
level in good forest on Mindanao (Heaney et al.,
1998; Heaney, 2001) and should also be sought
at lower elevations on Camiguin.

Two of the five specimens we captured were
females. An adult female taken on 17 March
1995, with enlarged mammae and weighing 71 g,
had recently given birth; it had three placental
scars (one left, two right), and the uterus was
swollen. The other female (62.5 g), probably
nulliparous, had small mammae. Two adult
males (60 g and 72 g) had scrotal testes (15 X
8 mm); the third male, taken on 24 May 1994,
was a small juvenile (28 g) with abdominal testes.
These scanty data imply that March, April, and
May are months of reproductive activity. Fe-
males have two pairs of mammae, indicating that
litter size is small.

Cranial and external measurements (Table 6)
are similar to but slightly larger than those from
Leyte and Mindanao (Kitanglad Range and Mt.
Apo; Rickart et al., 1993, 1998; Heaney et al.,
unpubl. data).

Specimens Examined — Total 5. Site 4 (2 msu-
iit); Site 5 (1 msu-iit); Site 7 (2 fmnh).

Rattus everetti (Gunther, 1879)

The common Philippine forest rat is a wide-
spread endemic species (excluding the Babuyan/
Batanes, Palawan, and Sulu groups) found in
primary and disturbed lowland, montane, and
mossy forest from sea level to 2400 m (Heaney et
al., 1991, 1998, 1999; Musser & Heaney, 1992;
Rickart et al., 1993).

During 1994-1995, we found this 250-350 g
rodent to be fairly common at all sampling sites
from disturbed lowland forest at 1000 m,
through montane forest to mossy forest at
1475 m; specimens from the 1960s were taken
at 500 m (Table 1, Fig. 1); on Mindanao, it
occurs down to sea level (Heaney, 2001; Heaney
et al., unpubl. data). It was trapped under root
tangles and beneath fallen logs, in hollow spaces
beneath live trees, and near mounds of soil and
rotting leaves.

Five parous adult females from May 1994 and
March 1995 had a mean weight of 271 ± 28.8
(range = 240-300 g, N = 5). During the same
months, five nulliparous females (with adult
pelage) weighed 188 ± 21.4 g (range = 160-
215 g), and four females with juvenile pelage
weighed 116 ± 31.0 g (range = 77-150 g).
Females have four pairs of mammae, indicating
moderate litter size. None were pregnant, and we
were unable to count placental scars; previous
studies also produced little data on litter size
(Heaney et al., 1999, unpubl. data). Males with
scrotal testes, recorded in May 1994 and March
1995, had a mean weight of 288 ± 69.8 g (range
= 210 375 g, N = 4), and the largest males had
testes size of 33 X 52 mm and 30 X 50 mm.
Three males with abdominal testes, during the
same months, weighed an average of 207 g
(range = 160-295 g).

Cranial and external measurements (Table 6)
are comparable to those from Biliran, Leyte, and
Maripipi but slightly smaller than series from
Dinagat, Mindanao (Kitanglad Range), and
Siargao (Heaney & Rabor, 1982; Heaney et al.,
1991, 1999, unpubl.; Rickart et al., 1993).

Specimens Examined— Total 28. Site 4 (6 msu-
iit); Site 5 (2 msu-iit); Site 6 (3 msu-iit); Site 7 (8

40

FIELDIANA: ZOOLOGY

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HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

41

fmnh); Site 8 (2 fmnh); Site 1 1 (4 dmnii); Site 12
(1 dmnh); Site 13 (2 dmnii).

Rutins exulans (Peale, 1848)

Known as the Polynesian rat or the spiny
rice-field rat, R. exulans is not native to the
Philippines; through mostly accidental dispersal
by humans, it occurs from Bangladesh to Easter
Island and throughout the Philippines, especially
as a pest in agricultural areas. On Camiguin, we
found this small (60-75 g) rat to be moderately
common in heavily disturbed lowland agricul-
tural land at ca. 150 m (Site 3) and in primary
montane forest at 1275 m (Site 7) but not at
other sites. Records from the 1960s also show it
to be present in mossy forest at ca. 1500 m. This
distribution is similar to that found on Negros,
where it occurs in high-density agricultural areas
and in mossy forest at an elevation of 1500-
1650 m (Heideman et al., 1987; Heaney et al.,
1989). Both Camiguin and Negros have very few
native rodents, and this may influence the ability
of nonnative species to invade the forest since the
nonnatives are absent or very rare in mature
forest on species-rich islands such as Leyte,
Luzon, and Mindanao (Heaney et al., 1989,
1999. unpubl. data; Rickart et al., 1993).

In March 1995, one nulliparous female weigh-
ing 41 g was trapped. Two adult males (71-76 g)
with scrotal testes and two young adults (49-
61 g) with scrotal testes were trapped at the same
time. They typically become reproductive at
a young age and have many large litters each
year (Barbehenn et al., 1973). Cranial and
external measurements (Table 6) are comparable
to those of Dinagat, Leyte, and Mindanao (Mt.
Kitanglad), but slight variations in most dimen-
sions are discernible (Heaney & Rabor, 1982;
Rickart et al., 1993; Heaney et al., unpubl. data).

Specimens Examined — Total 15. Site 3 (2 msu-
iit); Site 7 (5 fmnh); Site 12 (1 dmnh); Site 13 (7

DMNH).

Rattus tanezwni Temminck, 1844

Previously known in the Philippines as Rattus
rattus and Rattus mindanensis, the Oriental house
rat is a widespread nonnative rodent in the
Philippines; it occurs from Afghanistan to
Indomalaya. New Guinea, and Micronesia
(Musser & Carleton, 1993). This 140 200-g rat
is most often found as a pest in urban and
agricultural areas but can be common in
disturbed forest up to 1800 m (Heideman et al.,
1987; Heaney et al., 1989, 1991, 1998, unpubl.

data; Rickart et al., 1993). On Camiguin, we
found R. tanezwni to be very abundant in heavily
disturbed agricultural land at 150 m (Site 3;
Table 1). It was less common in secondary
lowland forest at 1000 m and in disturbed lower
montane forest at 1000 m (Site 4) and 1200 m
(Site 5), and a single individual was trapped in
primary mossy forest at 1400 m (Site 6). Four
nestling juveniles were found in a new slash-and-
burn clearing near Site 7, but no R. tanezwni
were caught in adjacent mature forest. Taken
together with records from the 1960s, these data
indicate that R. tanezwni occurs from sea level to
the highest peaks on Camiguin, although it tends
to be most common at lower elevations and in
disturbed habitats.

Pregnant females had a mean weight of 126 ±

57.6 g (range = 63-200 g, N = 4). Females with
large mammae but not pregnant had a mean
weight of 151 ± 16.7 g (range = 130-169 g, N =
9); three of these had placental scars (three, four,
and eight scars). Nulliparous females had a mean
weight of 93 ± 39.8 g (range = 67-160 g). Males
with scrotal testes had a mean weight of 169.4 ±

22.7 g (range = 141-199 g, N = 8); a subadult
with scrotal testes weighed 105 g. These data
indicate large litter size (up to eight) and very
early reproduction, as is typical for the species
(Barbehenn et al., 1973). External and cranial
measurements (Table 6) are comparable to those
of Biliran, Leyte, Maripipi, and Mindanao, with
only slight variation.

Specimens Examined — Total 69. Site 3 (21
msu-iit); Site 5 (4 msu-iit); Site 6 (1 msu-iit); Site
7 (4 fmnh); Site 9 (2 dmnh); Site 10 (6 dmnh);
Site 11 (3 dmnh); Site 12 (6 dmnh); Site 13 (22
dmnh).

Order Carnivora

Family Viverridae — Civets

Paradoxurus hermaphroditus Jourdan, 1837

The common palm civet occurs from Sri
Lanka to Hainan and the Lesser Sunda Islands
and is widespread in the Philippines (Heaney et
al., 1998). It has been recorded in agricultural
and forested areas from sea level up to at least
2400 m (Heaney et al., 1991, 1999, unpubl. data).
We did not capture any palm civets, but in
March 1995 we were given a mandible from
a specimen taken by local hunters in the
mountains above Mahinog, thus confirming the
earlier report by Gray (1843). A local hunter
identified P. hermaphroditus from pictures as
being common on Camiguin.

42

FIELDIANA: ZOOLOGY

Specimens Examined — Total 1.(1 fmnh).

Viverra tangalunga Gray, 1832

The Malay civet (or tangalung) is found from
peninsular Malaysia to Sulawesi and is wide-
spread in the Philippines (Heaney et al., 1998). It
has been found in primary and secondary
lowland, montane, and mossy forest from sea
level to at least 1700 m (Rickart et al., 1993;
Heaney et al., 1999). We did not obtain any on
Camiguin, but a local hunter identified Viverra
tangalunga from photographs as being present
on Camiguin; we tentatively accept this as a valid
record.

Order Artiodactyla
Family Suidae — Pigs

Sus philippensis Nehring, 1 886

The Philippine warty pig is a Philippine
endemic that occurs in the Greater Luzon,
Greater Mindanao, and Mindoro faunal regions;
its numbers are declining (Oliver, 1992, 1999). It
formerly was abundant from sea level to at least
2800 m in all habitats; now it is common only in
remote forests (Heaney et al., 1991, 1999,
unpubl. data; Oliver, 1992, 1999; Garcia &
Deocampo, 1997).

On Camiguin in March 1995, we observed
hoof marks of wild pigs from disturbed lowland
forest at 1 000 m up to primary montane forest at
1275 m elevation (Site 7). Near the sampling site
at 1475 m (Site 8), we saw an active pig nest, and
scattered pig trails were in clear evidence. A local
hunter said that they were commonly hunted and
often sold in a small local market at Owakan,
Mahinog Municipality, but not in coastal cities.
We were given two adult mandibles by hunters
from pigs captured in forest in the early 1 990s in
the mountains above Mahinog.

Specimens Examined — Total 2. Above Mahi-
nog (2 fmnh).

Analysis and Discussion

Adequacy of Sampling: What Is Present and What
Is Absent?

Before interpreting field data, it is necessary to
evaluate the extent to which they are complete
and reliable. In doing so here, we follow the
procedures used by Heaney et al. (1989, 1991,
1999) and Rickart et al. (1993).

Fruit Bats — Because pteropodid bats lack
sonar systems, they are easily captured in mist
nets. The 1960s surveys, which focused on birds,
yielded many fruit bat specimens (Heaney, 1984),
most of them almost certainly from mist nets.
Those efforts obtained six species of fruit bats
(Fig. 2). Our netting in 1992-1994 yielded no
additional species but was not extensive. We
consider it quite possible that some additional
species may be present, especially Eonycteris
spelaea and Rousettus amplexicaudatus, since
both occur very widely in the Philippines
(Heaney et al., 1998), and possibly the large
Acerodon jubatus and Pteropus vampyrus, which
fly high above the canopy and so are difficult to
capture. However, our sampling was sufficient
that we consider it quite unlikely that the species
that are abundant and easily captured on Mind-
anao and associated islands (Heaney et al., 1989,
unpubl. data; Rickart et al., 1993) are present on
Camiguin, especially Haplonycteris fischeri and
Ptenochirus minor and probably Megaerops
ecaudatus. Suitable habitat for the high-elevation
specialist Alionycteris paucidentata is very limited
in area on Camiguin, if present at all; although
we did limited netting in that habitat, we think it
very unlikely that this species is present.

Sampling along the elevation gradient proba-
bly has produced a partial picture of variation in
species richness (Fig. 2) but as noted may be
incomplete because our netting in 1992-1995 was
not extensive (Table 3).

Insectivorous Bats — Insectivorous bats use
sophisticated sonar systems to navigate. They are
thus difficult to capture in mist nets, which were
our only means of capturing them; further, our
mist-netting efforts in 1992-1995 were limited in
extent (see Heaney & Tabaranza, 2006a; Table 3).
For this reason, we believe that our sample of six
species is likely to be quite incomplete and is not
usable for estimates of species richness, as is
often the case (Heaney et al., 2002). Our data
contribute, however, to general knowledge of the
natural history of these poorly known animals.

Nonvolant Small Mammals — Shrews, ro-
dents, and other small mammals that can be
captured in traps were poorly sampled by the
1 960s field teams, which focused their efforts on
birds (and secondarily on fruit bats). During
1994 and 1995, we accumulated 2783 trap-nights
at six sites that included all major habitat types
on the island, yielding 188 captures (Table 1). A
species-accumulation curve for these data (Fig. 3)
shows that the number of species recorded for

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

No. Species

200 400 600 800 1000 1200 1400 1600 1800 2000 2200 2400 2600 2800

No. Trap-Nights

Fig. 3. Species accumulation curve for nonvolant small mammals on Camiguin Island. The points shown are
for Sites 1 and 3 to 6. as described in the text.

the island reached a peak of eight species
(including five nonnative species) by the end of
sampling at the second site (1055 trap-nights)
and showed no further gain as we approached
the total of nearly 2800 trap-nights. Local
hunters either were unfamiliar with squirrels or
knew them from Mindanao; given their conspic-
uousness (Heaney et al., unpubl. data), they are
very unlikely to be present. We conclude that the
island is unlikely to support other species of
small mammals that are present on Mindanao,
including gymnures (Podogymnura), tree shrews
(Urogale), squirrels, or murid rodents (e.g.,
Batomys, Limnomys, and Tarsomys). Further,
the sampling along the elevational gradients was
sufficiently intense that we are likely to have an
accurate assessment of the pattern of species
richness above about 800 m, although it must be
noted that extensive habitat destruction at lower
elevations may have had an extensive impact on
the current patterns (discussed below).

Medium and Large Mammals — We collected
information only on the presence and absence of
large mammals and a mixed set of medium-sized
species (none of which are readily captured in
traps such as the ones we used), not on detailed
habitat use. We observed carefully during all our
time in the field for any evidence of additional
species, and we interviewed many residents of the
islands and showed them photos of potential
species; some of these residents are active
hunters. Neither we nor our informants had
any evidence for tarsiers, flying lemurs, or deer
on the island, though many people knew of deer
and tarsiers on Mindanao, and a few knew of
flying squirrels and flying lemurs. We consider it
unlikely that species other than those listed here
are present given the voucher specimens we

obtained and the concordance between our
specimens and the information received from
local hunters. Thus, we conclude that deer are
absent, along with the smaller but conspicuous
species such as tarsiers, squirrels, and flying
lemurs.

Elevational Patterns

Though our limited netting prevents us from
commenting on the patterns of diversity and
abundance of bats along the elevational gradient
on Camiguin, our trapping data (Table 1) pro-
vide a reasonably complete picture of the general
patterns among nonvolant species. The highest
density we encountered was at Site 3, in
agricultural land at ca. 150 m. However, these
were all nonnative rats and a shrew, representing
only three species, a pattern that we have often
seen (e.g., Heaney et al., 1989; Rickart et al.,
1991). The five native small mammal species
were all present (or suspect to be present) from
1000 to 1300 m and declined in species richness
to two species near the peak of Mt. Timpoong at
Site 8 (Table 1). Density (measured as the
number captured per 100 trap-nights) more than
doubled from 1000 to 1275 m, then fell to a much
lower level near the peak at 1475 m. However,
much of the change in density was driven by the
remarkably large numbers of Suncus murium at
the upper elevations; they were not taken at 1000
or 1200 m (though they were taken at 150 m),
were fairly common at 1300 and 1475 m, and
were exceptionally abundant at 1275 m, where
we often saw and heard them during the day.
These patterns for species richness, though
limited in extent, are similar to those from
elsewhere in the Philippines, where species

44

FIELDIANA: ZOOLOGY

richness is generally low at low elevations,
increases to about the area of transition from
montane to mossy forest, and declines with
increasing elevation in mossy forest (Rickart et
al., 1991; Heaney, 2001). The pattern of overall
abundance is also similar to that documented
elsewhere, except that Suncus murinus is re-
markably abundant in the high-elevation mon-
tane and mossy forest, where usually it is absent
(Heaney, 2001). We have previously noted the
abundance of S. murinus in similar habitat only
once, on Negros Island (Heaney et al., 1989),
which is notably depauperate of small mammals
(Heaney, 1986). This raises a question: Is
Camiguin depauperate, relative to its area, in
comparison to other islands in the Philippines?
This question is addressed in the next section.

Biogeographic Implications

Bats — Because our focus in conducting field
studies concentrated primarily on nonvolant
mammals, we note only that we are confident
that the fruit bat community on Camiguin is at
least somewhat depauperate relative to similar
elevations on Mindanao and associated islands.
At least two species usually common at lower
and middle elevations, Haplonycteris fischeri and
Ptenochirus minor, are almost certainly absent.
Both of these are primarily associated with old-
growth rain forest and generally do not travel
long distances out from under the canopy of
forest (Heaney et al., 1989, 1998; Heideman &
Heaney, 1989; Rickart et al., 1993). Some
additional species probably also are absent
(Alionycteris paucidentata and Megaerops wet-
morei); they also are associated with old-growth
rain forest, though A. paucidentata in a distinctive
high-elevation habitat (M. wetmorei is poorly
known). This level of reduction in species
richness is striking in view of the tight correlation
between species richness and island area pre-
viously found (Heaney, 1991a). However, the
uncertainty regarding the potential absence of
Eonycteris and Rousettus leads us to be cautious
in defining the extent to which species richness is
low.

Nonvolant Mammals — The data are more
certain regarding both small mammals (shrews
and murid rodents) and the medium and larger
mammals (deer, squirrels, and so on). It seems
certain that many genera present in the lowlands
and in montane habitats on Mindanao and
associated islands are absent: tree shrews (Ur-

ogale), flying lemurs (Cynocephalus), tarsiers
(Tarsius), tree squirrels (Sundasciurus), pygmy
squirrels (Exilisciurus), flying squirrels (Peti-
nomys), a murid (Tarsomys echinatus), and deer
(Cervus). Among the murid rodents, Batomys is
also absent though often found in montane
forest on Mindanao and associated islands
(Rickart et al., 1993). Limnomys bryophilus, L.
sibuanus, and Tarsomys apoensis also occur on
Mindanao but only above about 1900 m eleva-
tion (Musser & Heaney, 1992; Heaney et al.,
1998; Rickart et al., 2003).

Those species of nonvolant mammals that are
present on Camiguin form a group that is quite
consistent in one respect. Not all lowland species
from Mindanao are present, but all those that
are present on Camiguin are among the relatively
few species that are common in the lowlands of
Mindanao — or were present before human de-
struction of lowland rain forests. As shown by
Musser and Heaney (1992), Heaney (2001), and
Heaney et al. (unpubl. data), Crocidura, Apomys,
Bullimus, Crunomys, Rattus everetti, Paradox-
urus, Viverra, and Sus are among the few
mammals that occur (or occurred until recently)
on Mindanao in lowland forest. With only one
exception (Tarsomys echinatus), all those low-
land species of Mindanao noted above as being
absent from Camiguin are arboreal species. In
other words, the nonvolant mammal fauna of
Camiguin is composed only of nonarboreal small
mammals from Mindanao (or are their sister
taxa) and all the lowland larger mammals (all of
which are also not arboreal). We conclude that
the mammalian fauna of Camiguin is highly
biased; that is, it is composed entirely of species
shared with and/or derived from Mindanao, but
it is not a random sample of the Mindanao
fauna; rather, it is comprised of species that
occur (or occurred before deforestation) on the
lowland forest floor of Mindanao, not in the
forest canopy and not in the montane or mossy
forest.

Does the absence of arboreal mammals mean
that Camiguin has a species-poor nonvolant
mammal fauna relative to other islands in the
Philippines? Perhaps surprisingly, the answer is
clearly no. With nine native nonvolant mam-
mals, Camiguin falls almost precisely on the
same species-area curve as the islands that made
up Greater Mindanao (Mindanao, Leyte, Bohol,
Biliran, and Maripipi). While the fauna is biased
toward lowland, ground-living murid rodents,
and small omnivorous carnivores, the species

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

45

richness is not reduced relative to islands that
were recently connected to Mindanao itself
(Heaney. 1986; Rickart et al., 1993). However,
the nonvolant small mammal community is
apparently sufficiently species-poor to allow
a nonnative shrew (Suncus minimis) to invade
primary montane forest. The presence of this
shrew in similar habitat has been noted on
Negros Island, which is also species poor
(Heaney et al., 1989), but not on other islands
in the Philippines (e.g., Rickart et al.. 1991.
1993). We predict that S. minimis will be found
in primary montane forest on other islands in the
Philippines with five or fewer native shrews and
rodents.

Conservation and Management

As noted by Heaney and Tabaranza (1997,
2006a), the native mammal fauna of Camiguin is
dependent on the continued survival of good-
quality forest at all elevations. The two mammal
species unique to Camiguin, in particular,
apparently depend on forest with little or no
disturbance, most of which currently occurs
in steep areas above 800 m elevation. These
forested areas are also crucial to the well-being
and stability of the human population, for
they are the source of water for the island and
protect the lowlands from potentially devastat-
ing floods and landslides. Additionally, our
observations indicate that the medium and large
mammals have been depleted by overhunting;
these require protection if they are to survive.
The people of Camiguin benefit both personally
and economically from the beautiful landscape
and seashores of the island, both through the
tourism they make possible and from the
environmental stability they engender. Protect-
ing the forests will benefit the people and wildlife
equally.

Acknowledgments

Our thanks go to the Protected Areas and
Wildlife Bureau of the Philippine Department of
Environment and Natural Resources, which
provided encouragement and permits, and the
Region 10 office of the DENR for logistical
support. Specimens for comparison were kindly
loaned by G. Hess (dmnh) and P. Myers (ummz)
and M. Carleton, L. Gordon, and H. Kafka

(lsnm). The manuscript was much improved by
comments from P. D. Heideman and E. A.
Rickart. We thank Lisa Kanellos for preparation
of the figures. Funding was provided by the
Marshall Field Fund. Ellen Thorne Smith Fund,
and the Barbara Brown Fund for Mammal
Research of the Field Museum of Natural
History.

Literature Cited

Balete, D. S.. B. R. Tabaranza, Jr., and L. R.
Heaney. 2006. An annotated checklist of the land
birds of Camiguin Island, Philippines, pp. 58-72. In
L. R. Heaney. ed.. The Mammals and Birds of
Camiguin Island. Philippines, a Distinctive Center
of Biodiversity. Fieldiana Zoology, n.s., 106: 1-72.

BARBEHENN, K.. J. P. SUMANGIL, AND J. B. LlBAY. 1973.

Rodents of the Philippine croplands. Philippine
Agriculturalist, 56: 217-242.

ESSELSTYN, J. A.. P. WlDMANN. AND L. R. HEANEY.

2004. The mammals of Palawan Island. Philippines.
Proceedings of the Biological Society of Washing-
ton. 117: 271-302.

Fooden. J. 1995. Systematic review of Southeast Asian
longtail macaques. Macaca fascicularis (Raffles.
[1821]). Fieldiana Zoology, n.s., 81: 1 206.

Garcia. M. G. P., and A. H. Deocampo. 1997. Wild
pig meat trading in Iligan City, Mindanao. Sylva-
trop. 5(1995):29-34.

Goodman. S. M., and N. R. Ingle. 1993. Sibuyan
Island in the Philippines-threatened and in need of
conservation. Oryx, 27: 174-180.

Gray, J. E. 1843. List of the specimens of mammals in
the collection of the British Museum. British
Museum (Natural History).

Hall, R. 1998. The plate tectonics of Cenozoic SE
Asia and the distribution of land and sea, pp. 99-
132. In R. Hall and J. D. Holloway, eds.. Bio-
geography and Geological Evolution of SE Asia.
Backhuys Publishers. Leiden.

— . 2002. Cenozoic geological and plate tectonic
evolution of the SE Asia and the SW Pacific:
Computer-based reconstructions, model and anima-
tions. Journal of Asian Earth Sciences, 20: 353^31.

Heaney, L. R. 1984. Mammals from Camiguin Island,
Philippines. Proceedings of the Biological Society of
Washington, 97: 119-f25.

— . 1986. Biogeography of mammals in Southeast
Asia: Estimates of rates of colonization, extinction,
and speciation. Biological Journal of the Linnean
Society. 28: 127-165.

1991a. An analysis of patterns of distribution

and species richness among Philippine fruit bats
(Pteropodidae). Bulletin of the American Museum
of Natural History. 206: 145-167.

1991b. A synopsis of climatic and vegetational

change in Southeast Asia. Climatic Change, 19: 53-61.

46

FIELDIANA: ZOOLOGY

. 1993. Biodiversity patterns and the conserva-
tion of mammals in the Philippines. Asia Life
Sciences, 2: 261-274.

2000. Dynamic disequilibrium: A long-term,

large-scale perspective on the equilibrium model of
island biogeography. Global Ecology and Biogeog-
raphy, 9: 59-74.

2001. Small mammal diversity along eleva-

of Camiguin Island, Philippines, a Distinctive Cen-
ter of Biodiversity, Fieldiana Zoology, n.s., 106:
1-72.

2006b. A new species of forest mouse, genus

tional gradients in the Philippines: An assessment of
patterns and hypotheses. Global Ecology and Bio-
geography, 10: 15-39.

2004. Conservation biogeography in an

oceanic archipelago, pp. 345-368. In M. V. Lomo-
lino and L. R. Heaney, eds., Frontiers of Bio-
geography, New Directions in the Geography of
Nature. Sinauer Associates, Sunderland.
Heaney, L. R., D. S. Balete, L. Dolar, A. C. Alcala,

A. Dans, P. C. Gonzales, N. R. Ingle, M. Lepiten,
W. L. R. Oliver, P. S. Ong, E. A. Rickart, B. R.
Tabaranza, Jr., and R. C. B. Utzurrum. 1998. A
synopsis of the mammalian fauna of the Philippine
Islands. Fieldiana Zoology, n.s., 88: 1-61.

Heaney, L. R., D. S. Balete, E. A. Rickart, R. C. B.
Utzurrum, and P. C. Gonzales. 1999. Mammalian
diversity on Mt. Isarog, a threatened center of
endemism on southern Luzon Island, Philippines.
Fieldiana Zoology, n.s., 95: 1-62.

Heaney, L. R., P. C. Gonzales, R. C. B. Utzurrum,
and E. A. Rickart. 1991. The mammals of
Catanduanes Island: Implications for the biogeog-
raphy of small land-bridge islands in the Philippines.
Proceedings of the Biological Society of Washing-
ton, 104: 399^115.

Heaney, L. R., P. D. Heideman, E. A. Rickart, R. C.

B. Utzurrum, and J. S. H. Klompen. 1989.
Elevational zonation of mammals in the central
Philippines. Journal of Tropical Ecology, 5:
259-280.

Heaney, L. R., and D. S. Rabor. 1982. Mammals of
Dinagat and Siargao Islands, Philippines. Occasion-
al Papers of the Museum of Zoology, University of
Michigan, 699: 1-30.

Heaney, L. R., and J. C. Regalado, Jr. 1998.
Vanishing Treasures of the Philippine Rain Forest.
The Field Museum, Chicago.

Heaney, L. R., and E. A. Rickart. 1990. Correlations
of clades and clines: Geographic, elevational, and
phylogenetic distribution patterns among Philippine
mammals, pp. 321-332. In G. Peters and R.
Hutterer, eds., Vertebrates in the Tropics. Museum
Alexander Koenig, Bonn.

Heaney, L. R., and M. Ruedi. 1994. A preliminary
analysis of biogeography and phylogeny of Croci-
dura from the Philippines. Special Publication
Carnegie Museum of Natural History, 18: 357-
377.

Heaney, L. R., and B. R. Tabaranza, Jr. 1997. A
preliminary report on mammalian diversity and
conservation status of Camiguin Island, Philippines.
Sylvatrop (1995), 5: 57-64.

. 2006a. Introduction to the mammal and land

bird faunas of Camiguin Island, Philippines, pp. 1-
13. In L. R. Heaney, ed., The Mammals and Birds

Apomys (Rodentia, Muridae) from Camiguin
Island, Philippines, pp. 14-27. In L. R. Heaney,
ed., The Mammals and Birds of Camiguin Island,
Philippines, a Distinctive Center of Biodiversity,
Fieldiana Zoology, n.s., 106: 1-72.

Heaney, L. R., E. K. Walker, B. R. Tabaranza, Jr.,
and N. R. Ingle. 2002. Mammalian diversity in the
Philippines: An assessment of the adequacy of
current data. Sylvatrop, 10(2000):6-27.

Heideman, P. D. 1989. Delayed development in
Fischer's pygmy fruit bat, Haplonycteris fischeri, in
the Philippines. Journal of Reproduction and
Fertility, 85: 363-382.

. 1995. Synchrony and seasonality of reproduc-
tion in tropical bats. Symposium of the Zoological
Society of London, 67: 151-165.

Heideman, P. D., J. A. Cummings, and L. R. Heaney.
1993. Reproductive timing and early embryonic
development in an Old World fruit bat, Olopteropus
cartilagonodus (Megachiroptera). Journal of Mam-
malogy, 74: 621-630.

Heideman, P. D., and L. R. Heaney. 1989. Popu-
lation biology and estimates of abundance of
fruit bats (Pteropodidae) in Philippine submontane
rainforest. Journal of Zoology, London, 218:
565-586.

Heideman, P. D., L. R. Heaney, R. T. Thomas, and
K. R. Erickson. 1987. Patterns of faunal diversity
and species abundance of non-volant small mam-
mals on Negros Island, Philippines. Journal of
Mammalogy, 68: 884-888.

Heideman, P. D., and K. S. Powell. 1998. Age-
specific reproductive strategies and delayed embry-
onic development in an Old World fruit bat,
Ptenochirus jagori. Journal of Mammalogy, 79:
295-311.

Ingle, N. R. 1992. The natural history of bats on Mt.
Makiling, Luzon Island, Philippines. Silliman Jour-
nal, 36: 1-26.

Ingle, N. R., and L. R. Heaney. 1992. A key to the
bats of the Philippine Islands. Fieldiana Zoology,
n.s., 69: 1-44.

Klingener, D., and G. K. Creighton. 1984. On small
bats of the genus Pteropus from the Philippines.
Proceedings of the Biological Society of Washing-
ton, 97: 395-403.

Koopman, K. F. 1989. Distributional patterns of Indo-
Malaysian bats (Mammalia: Chiroptera). American
Museum Novitates, 2942: 1-19.

Lepiten, M. 1997. The mammals of Siquijor Island,
central Philippines. Sylvatrop, 5(1 995): 1-1 7.

MlTTERMEIER, R. A., P. R. GlL, AND C. G. MlTTERMEIER,

eds. 1997. Megadiversity: Earth's Biologically Wealth-
iest Nations. CEMEX, Monterrey, Mexico.

MlTTERMEIER, R. A., N. MYERS, P. R. GlL, AND C. G.

Mittermeier, eds. 1999. Hotspots, Earth's Biolog-
ically Richest and Most Endangered Terrestrial
Ecosystems. CEMEX, Mexico City.

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND

47

Musser, G. G. 1982. Results of the Arehbold
Expeditions, No. 108. The definition of Apomys,
a native rat o( the Philippine Islands. American
Museum Novitates, 2746: 1 43.

MrssiR, G. G., and M. D. Carleton. 1993. Family
Muridae. pp. 501 755. /// Wilson. D. and
D. M. Reeder, eds.. Mammal Species of the World:
A Taxonomic and Geographic Reference, 2nd
ed. Smithsonian Institution Press, Washington,
DC.

Mi ssi r, G. G., and L. R. Heaney. 1992. Philippine
rodents: Definitions of Tarsomys and Limnomys
plus a preliminary assessment of phylogenetic
patters among native Philippine murines (Murinae.
Muridae). Bulletin of the American Museum of
Natural History, 211: 1-138.

Mi ssi r, G. G., L. R. Heaney, and B. R. Tabaranza,
Jr. 1998. Philippine rodents: Redefinition of known
species of Batomys (Muridae, Murinae) and descrip-
tion of a new species from Dinagat Island. Philippines.
American Museum Novitates. 3237: 151.

Oliver, W. L. R. 1992. The taxonomy, distribution,
and status of Philippine wild pics. Silliman Journal.
36: 55 64.

— . 1999. Taxonomy and conservation of Philip-
pine wild pigs An update. Sylvatrop. 8(1998):27-
41.

Peterson, R. L., and M. B. Fenton. 1970. Variation
in bats of the genus Harpyionycteris, with the
description of a new race. Royal Ontario Museum
Life Sciences Occasional Papers. 17: 115.

Rickart, E. A., L. R. Heaney, D. S. Balete, and B.
R. Tabaranza, Jr. 1998. A review of the genera
Crunomys and Archboldomys (Rodentia. Muridae,

Murinae) with descriptions of two new species
from the Philippines. Fieldiana Zoology, n.s.. 89:
1 24.

Ric kart, E. A.. L. R. Heaney, P. D. Heideman. and
R. C. B. Utzurrum. 1993. The distribution and
ecology of mammals on Leyte, Biliran, and Maripipi
Islands, Philippines. Fieldiana Zoology, n.s., 72:
1 62.

Rickart, E. A., L. R. Heaney, and B. R. Tabaranza,
Jr. 2002. Review of the Biillimus (Muridae: Mur-
inae) and description of a new species from
Camiguin Island, Philippines. Journal of Mammal-
ogy, 83: 421^436.

— . 2003. A new species of Limnomys (Rodentia:
Muridae: Murinae) from Mindanao Island. Philip-
pines. Journal of Mammalogy. 84: 1443-1455.

Rickart, E. A., L. R. Heaney, and R. B. Utzurrum.
1991. Distribution and ecology of small mam-
mals along an elevational transect in southeastern
Luzon. Philippines. Journal of Mammalogy. 72:
458-469.

Ruedas, L. A. 1994. Morphological and ecological
variation in Otopteropus cartilagonodus Kock 1969
(Mammalia: Chiroptera; Pteropodidae) from Lu-
zon, Philippines. Proceedings of the Biological
Society of Washington, 107: 1-16.

Steppan, S., C. Zawadski, and L. R. Heaney. 2003. A
molecular assessment of phylogenetic relationships
and patterns of phylogenesis in the Philippine murid
rodent Apomys. Biological Journal of the Linnean
Society, 83: 699-715.

Utzurrum. R. C. B. 1992. Conservation status of
Philippine fruit bats (Pteropodidae). Silliman Jour-
nal, 36: 27-46.

48

FIELDIANA: ZOOLOGY

A New Species of Hanging-Parrot (Aves: Psittacidae:
Loriculus) from Camiguin Island, Philippines

Jose G. Tello,1'2'3 Jacob F. Degner,1 John M. Bates,1 and David E. Willard1

Abstract

A new species of Hanging-Parrot or Colasisi, Loriculus, is described from a series of 23
specimens (19 males, 4 females) collected in the 1960s on Camiguin Island, Camiguin
Province, Philippines, at elevations between 300 and 1350 m. The new species lacks sexual
dimorphism in plumage coloration, which distinguishes it from all other members of the L.
philippensis group and all other Loriculus. The overall color pattern of the new species appears
most like females of L. p. worcesteri and L. p. apicalis but differs in plumage characteristics
(the width and extension of the orange-scarlet crown patch, the amount and intensity of blue
in the face and thighs, and the intensity of the blue in the tail above inner edges and the tail
below). In addition, males of the new species are larger than males of nearby populations of L.
philippensis, having significantly longer tails and wing chords. Nothing is known about the
habits of the new species; however, the small size of the island of Camiguin, coupled with
extensive deforestation, makes the status of the new species a significant conservation
concern.

Introduction

The Philippine Hanging-Parrot or Colasisi
{Loriculus philippensis) has ten described sub-
species distributed throughout the islands of the
Philippines (Dickinson et al., 1991; Collar, 1997;
Juniper & Parr, 1998; Kennedy et al., 2000). The
subspecies L. p. apicalis has been reported to
occur on the islands of Bazol, Balut, Camiguin,
Dinagat, Mindanao, and Siargao (Fig. 1). How-
ever, Austin Rand, a former Field Museum

1 Field Museum of Natural History, 1400 South
Lake Shore Drive, Chicago, IL 60605-2496, U.S.A.

2 Biological Sciences, University of Illinois at
Chicago, Chicago, IL 60607, U.S.A.

3 Current address: Division of Vertebrate Zoology-
Ornithology, American Museum of Natural History,
Central Park West at 79th Street, New York, NY
10024-5192, U.S.A.

curator and Philippine expert (e.g., Rand &
Rabor 1960, 1969), had penciled the notation
"subsp. nov" on the tag of a specimen from
Camiguin in the fmnh collection. He never
published a description. Here, we quantitatively
evaluate the external morphology and compare
the plumage color of specimens referred to L. p.
apicalis from Camiguin with L. p. apicalis from
Mindanao and specimens of other subspecies of
L philippensis. Our results demonstrate that the
Camiguin population of L. p. apicalis is separa-
ble from all other populations of L. philippensis
in plumage. It is further separable from all
neighboring populations in body size. We argue
that these differences warrant designating this
population as a distinct species and not a sub-
species of L. philippensis. Similar arguments have
been made in separating L. bonapartei of the
Sulu Archipelago from the L. philippensis com-
plex (Juniper & Parr 1998). We present a formal

FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, PP. 49-57

49

Fig. 1. Map of the central and southern Philippines showing the locations of islands referred to in the text.

50

FIELDIANA: ZOOLOGY

description of the new taxon based on plumage
and morphology of adult males.

Methods

All qualitative color comparisons were made
under natural light. Color names follow Smithe
(1975), and each color name (capitalized) is
followed by its number in parentheses. J.F.D.
measured wing chord, tarsus length, tail length
(from point of insertion of central rectrices to tip
of longest rectrix), culmen length, bill length
(from anterior edge of nostril), bill height (at
anterior edge of nostril), bill width (at anterior
edge of nostril), and gape width with calipers to
the nearest 0.05 mm. Specimens were measured
randomly to avoid any investigator bias.

All statistical analyses were carried out using
the program Statistica (Statsoft Inc. 1995).
Mensural data were tested for normality using
Kolmogorov-Smirnov tests and Lilliefors prob-
abilities prior to all the analyses. Mensural
differences between males and females within
each study population were evaluated using one-
way analyses of variance (ANOVAs). We used
both univariate (one-way ANOVAs and Bonfer-
roni post hoc tests) and multivariate analysis
(principal components and discriminant func-
tion) on log-transformed data to test for
mensural differences between specimens from
Camiguin and those from neighboring popula-
tions of L. philippensis. Multivariate analyses
were used to reduce the dimensionality of data
and facilitate the analysis of morphology in two
or three dimensions (Pimentel, 1979); we used the
varimax raw method to rotate the three compo-
nents that are reported in the principal compo-
nents analysis in order to improve interpretabil-
ity of the resulting patterns. Collecting localities
are described by Heaney and Tabaranza (2006a).

Loriculus camiguinensis, new species

Camiguin Hanging-Parrot

Holotype — Field Museum of Natural History
No. 284389, adult male from Kasangsangan,
Municipality of Catarman, Camiguin Province,
Camiguin Island, Philippines, elevation between
1000 and 2000 ft (300-600 m; approx. 9°11'N,
124°40'E; see Heaney & Tabaranza, 2006a, for
more explanation of this and other collecting

localities on Camiguin Island), 18 June 1968,
collected by D. S. Rabor and W. Sanguila.

Diagnosis — A Loriculus hanging-parrot with
characteristics of the philippensis group (see
Front Plate). In contrast to other members of
this group, L. camiguinensis is characterized by
a lack of sexual dimorphism in plumage color-
ation. The overall color pattern of L. camigui-
nensis is most like females of L. p. worcesteri
from Bohol, Leyte, and Samar and L. p. apicalis
from Mindanao but differs as follows: (1) the
scarlet of the crown of L. camiguinensis does not
extend as far onto the bright olive green nape as
it does in both males and females of L. p. apicalis
and L. p. worcesteri; this pattern differs from that
of chrysonotus, siquijorensis, regulus, bournsi,
mindorensis, and philippensis, in which the scarlet
crown is highly reduced. (2) The width of the
scarlet crown in L. camiguinensis narrows at the
rear edge instead of being rounded as in all other
populations. (3) The scarlet (sometimes orange)
throat patch that is typical of males in L.
philippensis is lacking in L. camiguinensis; five
of the Camiguin males had data on gonadal
development, reporting enlarged or slightly
enlarged testicles, which gives an indication of
their adult condition. (4) The face of L.
camiguinensis is extensively turquoise blue and
differs from that of females of L. philippensis
subspecies in that the blue of the face is darker
and more extensive, extending over the eye and
onto the throat. (5) The turquoise blue in the
thighs of L. camiguinensis is darker than that
of females of L. philippensis populations. (6)
The blue in the inner edges of the rectrices above
and throughout below is darker in L. camigui-
nensis. (7) Mean wing chord and tail length of
males and tail length of females of L. camigui-
nensis are significantly longer than those of
nearby L. philippensis subspecies (Tables 1 and
2). (8) The overall green plumage is a darker
shade with less of a yellowish tinge, especially on
the back.

Description of Holotype — General plumage
Parrot Green (160) with slightly orange tinge in
the upperparts, more yellowish tinge on under-
pays; forehead and forepart of crown Scarlet
(14) fading to orange at rear-edge; thin Orange
Yellow (18) band on nape; lores, chin, cheek, and
throat closest to Turquoise Blue (65); rump and
upper tail-coverts scarlet; Turquoise Green (64)
markings on the sides of the rump; thighs slightly
paler Turquoise Blue; rectrices Emerald Dark
Green (262) above with dark Cerulean Blue (67)

TELLO ET AL.: A NEW SPECIES OF HANGING-PARROT

51

tinges on inner edges of all but central reetrices;
rectrices dark Cerulean Blue below; flight feath-
ers black above with Emerald Dark Green outer
edges, flight feathers black below with inner
edges with extensive Cerulean Blue; greater
underwing-coverts Cerulean Blue and lesser un-
derwing-coverts closest to Spectrum Green (62).
Soft part colors of dried specimen: upper
mandible closest to Spectrum Orange (17) at
base grading to yellow with Gray Horn (91) at
tip and along tomia; lower mandible with similar
pattern, but orange at the base; cere Grayish-
Horn; feet and tarsi, yellow-horn.

Measurements of Holotype (mm) — Wing
(99.8), tarsus (11.7), tail (49.5), culmen (18.0),
bill length (15.0), bill height (11.1), bill width
(6.4), gape width (8.2).

Distribution — Loriculus camiguinensis is
known only from the forests of Camiguin Island.
Specimens have been collected between 1000 and
4500 ft (300-1350 m) in the municipalities of
Catarman and Mahinog (Balete et al., 2006;
Heaney & Tabaranza, 2006a).

Etymology — We name this species after the
Philippine Island of Camiguin, to which this
species appears to be endemic.

Specimens Examined — We examined the fol-
lowing specimens from Field Museum of Natural
History (fmnh) and Delaware Museum of
Natural History (dmnh):

L. camiguinensis (Camiguin Island) (19 males,
4 females, all known specimens of this new
taxon): Camiguin Province: Catarman Munici-
pality; Kasangsangan (males: fmnh holotype,
fmnh 284391, 284392, 284393; dmnh 19950,
19958, 19960, 19961, 19962: females fmnh
284390; dmnh 19959); Camiguin Province: Cat-
arman Municipality; Catarman Mountain (male:
dmnh 19949); Camiguin Province: Mahinog
Municipality; Matugnao, Mt. Timpoong (males:
fmnh 286742, 286743, 286744, 286745; dmnh
19951, 19952, 19953, 19954, 19965; females:
fmnh 286746, 286747).

L. p. apicalis (Mindanao Island) (19 males, 5
females): North Cotabato Province: Mt. Apo,
Todaya (male: fmnh 184090); North Cotabato
Province: Mt. Apo, Galog (male: dmnh 36227);
Agusan del Norte: Mt. Hilong-Hilong, Lewed
(male: fmnh 275003); Misamis Occidental: Zam-
boanga Peninsula, Mt. Malindang, Gandwan
(males: fmnh 227136, 227138, 227139; female:
fmnh 227137); Misamis Occidental: Zamboanga
Peninsula. Mt. Malindang, Masawan (males:
fmnh 227134, 227135); Davao Oriental Province;

Mati: Mt. Mayo, Limot (male: fmnh 277864);
Misamis Oriental Province: Manticao: Tuod,
Camp Dundue (male: fmnh 283788; female
fmnh 283787); Misamis Oriental Province: Opol:
Malubato (male: fmnh 283785); South Cotabato:
Tupi: Mt. Matutum (male: fmnh 279330);
Bukidnon Province: Malaybalay, Mt. Katanglad
(male: fmnh 262475, 262476; female: fmnh
262474); Bukidnon Province: Lantapan: Ko-
toon, Mt. Katanglad SE slope (male: dmnh
2983); Lanao Norte Province: Iligan City,
Mainit, Mahayahay (female: fmnh 283786);
Surigao del Sur Province: Car-Can-Mad-Lan
area (female: fmnh 275002); Zamboanga del Sur
Province: Zamboanga (male: dmnh 36993);
Davao Oriental Province: Sigaboy (males: dmnh
36224, 36226); Davao del Sur Province: Padada
(male: dmnh 36233).

L. p. worcesteri (11 males, 8 females): Bohol
Island: Bohol Province: Sierra Bullones (males:
fmnh 223025, 223026, 223029, 223030, 223034,
223036, 223037; females: fmnh 223027, 223327,
223028, 223033, 223035, 223039); Leyte Island:
Leyte Province: Burauen, Buri, Ma-Alngon
(male: fmnh 276302; female: fmnh 276300);
Leyte Province: Burauen, Buri, Mt. Lobi range,
Tambis (male: fmnh 276299; female: fmnh
276298); Samar Province: Mt. Capotoan (male:
fmnh 247411); Western Samar Province: Matu-
guinao (male: fmnh 247410).

L. p. regulus (Negros Island) (5 males, 4
females): Negros Oriental Province: Bayawan,
Basay (male: fmnh 257121); Negros Oriental
Province: Santa Catalina, Inubungan (male:
fmnh 219314; male: fmnh 188579); Negros
Oriental Province: Sicopon River (male: fmnh
185483); Negros Oriental Province: Amio (males:
fmnh 188545, 188548; females: fmnh 188544,
188553); Negros Oriental Province: Pamo-at
(male: fmnh 188550).

L. p. chrysonotus (captive specimen, presum-
ably from Cebu Island) (1 male): Cebu Province:
Exact locality unknown (fmnh 252666).

L. p. siquijorensis (Siquijor Island) (1 male):
Siquijor Province: San Juan: Tag-ibo (fmnh
222741).

L. p. bournsi (Sibuyan Island) (1 male, 1
female): Romblon Province: Goangan, 3 km SE
Magdiwang (male: fmnh 358288); Romblon
Province: Exact locality unknown (female: fmnh
11081).

L. p. mindorensis (Mindoro Island) (1 male,
1 female): Oriental Mindoro Province:
Calapan (male: fmnh 19927); Occidental Mind-

52

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TELLO ET AL.: A NEW SPECIES OF HANGING-PARROT

53

Table 2. Univariate statistical comparisons be-
tween /.. camiguinensis and adjacent populations ol' L.
philippensis. Populations ( 1 = L. camiguinensis: 2 = L.
p. apicalis; 3 = L. p. worcesteri; and 4 = L p. regains)
are ordered based on their mean variation, from
smallest to largest (left to right). * F values signifi-
cantly different at P < 0.05 (one-way analysis oi'
variance and Bonferroni post-hoc test). For each
morphometric variable, populations united by the
underlines showed non-significant differences.

Morphometric
variables

Populations

Wing chord

30.5*

Tail length

23.1*

Tarsus length

14.8*

Gape width

9.8*

Bill length

3.7*

Total culmen

3.5*

Bill height

4.0*

Bill width

5 2*

2

3

4

2

4

3

2

3

4

2

3

4

2

4

3

2

3

4

4

2

3

2

4

3

oro Province: Abra de Hog (female: fmnh
210845).

L. p. philippensis (Luzon Island) (1 male. 1
female): Bataan Province: Mariveles (male: fmnh
73966); Cagayan Province: Sierra Madre, Mt.
Cagua (female: fmnh 258827).

Morphometric Differences — One-way ANO-
VAs on all known L. camiguinensis specimens
and specimens in the fmnh collection from
neighboring islands showed that certain variables
differed between sexes in some of the study
populations, but no consistent pattern was found
(Table 1). Because of the small sample of females
available for this study, we include only males in
the analyses.

Results of the one-way ANOVAs (effect =
population) and Bonferroni post hoc tests
showed that males of L. camiguinensis are
morphologically distinct from males of other L.
philippensis populations. Wing chord, tail length,
and tarsus length were significantly longer in L.
camiguinensis males than in the other popula-
tions (Table 2).

Principal components analysis for males from
all populations resulted in three rotated (varimax
raw) factors with Eigenvalues close to or greater
than 1.0, which together explained 73.2% of the
total variance (Table 3). Axes defined by the first
and third component (particularly the third)
demonstrate separation of L. camiguinensis from
the other populations (Fig. 2). The first compo-
nent explained 45.2c/c of the total variance and

Table 3. Fraction of total variance explained by
each of the first three components of the principal
components analyses. The three components were
rotated (see Methods).

Morphometric
variables

PCI

PCII PCIII

Wine chord 0.25 0.29 0.84

Tarsus length 0.29 0.31 0.40

Tail length 0.16 0.04 0.94

Total culmen 0.14 0.88 0.18

Bill length 0.05 0.92 0.09

Bill height 0.77 0.15 0.17

Bill width 0.80 -0.04 0.21

Gap width 0.71 0.33 0.35

Eigenvalue 3.62 1.34 0.90

Explained variance (%) 45.21 16.79 11.21

had high positive correlations with bill width, bill
height and gape width (0.80, 0.77, and 0.71.
respectively). The second component explained
16.8% of the total variance and had high positive
correlation with bill length and culmen length
(0.92 and 0.88, respectively). The third compo-
nent explained a further 1 1 .2% of the variance
and had the highest positive correlation values
for tail length and wing chord (0.94 and 0.84.
respectively). It was this latter axis that differ-
entiates L. camiguinensis from the other popula-
tions.

A discriminant function analysis using these
morphometric variables was also significant (P
< 0.001) and correctly classified 88% of the cases
(plot not shown). The standardized coefficients
of the discriminant function separating males of
L. camiguinensis from those of other L. philip-
pensis populations weighted tarsus length and
wing chord heavily (0.80 and 0.80, respectively),
followed by tail length (0.46) and bill length
(0.40), with all the other variables having
coefficients under 0.32.

A Question of Correctly Sexed Specimens —
Unpublished concern has been expressed re-
garding the ability of Rabor and his field
assistants to accurately determine the sex of
specimens. This possibility presents a serious
issue with respect to interpreting the data at
hand. In an attempt to verify sexing. we tried
unsuccessfully to amplify DNA from toe pads of
some specimens using commercially available
primers for sex-linked DNA. However, for the
following reasons, we remain convinced that
there is good reason to believe that the issue
of sexing does not overshadow the validity of
this taxon. It is our experience that mis-sexing

54

FIELDIANA: ZOOLOGY

o

L camiguinensis
L p. apicalis
L p. worcesteh
L p. regulus

Fig. 2. Results of the Principal Components Analysis of morphological data from populations of Philippine
Loriculus, on the first three axes; see text for details.

O

o

A

most commonly involves undeveloped gonads,
and there are clearly labeled tags indicating
that the gonads of some specimens were de-
veloped. If mistakes were made, the plumage
characters of the specimens described above
would logically argue that all males were
misidentified as females. Again, this seems highly
unlikely given the number of males (19).
Furthermore, it seems highly unlikely that
a series of 23 randomly collected parrots would
all have been females. Certainly, there is nothing
like this in other series of Loriculus collected by
Rabor in other parts of the Philippines. Thus,
while we cannot say that all specimens are
unequivocally identified to sex correctly, we feel
that adults of both males and females are
included in these series.

Discussion

Our results demonstrate that L. camiguinensis
is diagnosable from populations of L. philippen-
sis in plumage. It also differs in size from all

neighboring populations. We were able to make
direct comparisons with specimens of all sub-
species except L. p. dohertyi (Basilan). It is
possible that L. camiguinensis is more closely
related to some parts of the L. philippensis group
than others, which would make L. philippensis
paraphyletic (Funk & Omland 2003). Despite
this possible relationship to the widespread L.
philippensis group, we believe L. camiguinensis
sufficiently differentiated to be beyond concerns
expressed about recognizing new species based
on minor morphologic differences (e.g., Collar et
al., 1999). Based on geographic distance and the
overall pattern of plumage coloration, L. cami-
guinensis most closely resembles populations of
L. p. apicalis and L. p. worcesteri, but no
phylogenetic analyses exist yet for these taxa.
The comparatively dull plumage of the male
of L. camiguinensis is consistent with the
documented tendency for some insular bird
populations to lose bright plumage, leading to
a lack of sexual dichromatism (see references in
Peterson, 1996); L. camiguinensis is the only
member of the genus without sexual dichroma-
tism in plumage.

TELLO ET AL.: A NEW SPECIES OF HANGING-PARROT

The recognition of this distinctive taxon
coincides with recent surveys of the small
mammal fauna of Camiguin Island that have
discovered two new species of rodents (Rickart et
al., 2002; Heaney et al., 2006). Camiguin is
believed to be the smallest Philippine island to
harbor endemic species of birds and mammals
(Heaney & Tabaranza, 1997, 2006a). The island
has been continuously isolated from its large
southern neighbor, Mindanao, even during
periods of low sea level during the "ice ages"
of the Pleistocene, when sea levels dropped to
120 m below present levels (Heaney, 1986,
1991a, 1991b; Fairbanks, 1989; Heaney and
Tabaranza, 1997, 2005b; Heaney and Regalado,
1998; Hanebuth et al., 2000), and this may have
played a role in the differentiation of Camiguin's
fauna from that of Mindanao (Steppan et al.,
2003).

The value of museum collections is well
illustrated with this description. These collec-
tions were essential in the recognition and
documentation of L. camiguinensis. Had there
not been a series of specimens available for
study, we would have likely dismissed differences
in the new taxon as possibly aberrant or
immature plumage or an error in sexing of
a specimen (a female incorrectly identified as
a male; see above). However, the presence of
a series of specimens from different localities
(with data on gonadal development) has allowed
us to compile meaningful data sets on morpho-
logical variation and assess within-population
variation in color.

This new species also illustrates the need for
additional taxonomic and systematic research on
the Loriculus hanging-parrots to understand the
evolutionary patterns in the group and to
evaluate the possibility that some of the other
allopatric forms of L philippensis may also
deserve species status. The issue of assessing the
taxonomic status of allopatric populations in the
Philippines has long been recognized as a chal-
lenge for conservationists (Collar et al., 1999;
Peterson et al., 2000). To date, little attention has
been given to the conservation plight of Loriculus
parrots. For instance, none was included by
Collar et al. (1999) in their list of threatened
Philippine bird species. This lack of attention can
be directly correlated with the designation of L.
philippensis as a polytypic species. The plight of
these populations is cause for concern, as
Loriculus p. chrysonotus from the island of Cebu
is believed to be extinct (Forshaw, 1989; Mallari

et al.. 2001) and another form, L. p. siquijorensis,
may be extinct as well (Forshaw, 1989; Kennedy
et al., 2000). The combined populations of
Mindoro, Sibuyan, Negros, Panay, Tablas,
Romblon, Masbate, Ticao, Guimaras, and Basi-
lan (including mindorensis, bournsi, regulus, and
dohertyi) may total no more than 5000 individ-
uals (Juniper & Parr, 1998). The current popu-
lation size of L. camiguinensis is not known (but
see Heaney and Tabaranza, 2006a, for an
assessment of remaining habitat on the island).
Without field data on its status, we defer from
suggesting how this species should be character-
ized under international threat criteria (IUCN
Species Survival Commission, 1994). However,
because Camiguin is a small island that has
experienced extensive deforestation, the conser-
vation status of this newly described species
clearly requires assessment. Field study is needed
to establish the population size and requirements
as a prerequisite for conservation planning and
action.

Acknowledgments

We are grateful to G. Hess of the Delaware
Museum of Natural History for loan of speci-
mens under his care and to M. Skakuj for
painting the plate. Bruce Patterson and P.
Wagner provided help with data analyses, and
S. Bober helped in preparing Figure 1. The
manuscript was greatly improved by comments
from D. S. Balete, A. Carnaval, L. R. Heaney, S.
Ware, P. C. Rasmussen, and N. J. Collar,
although this does not imply that all agree with
the status we convey on this taxon.

Literature Cited

Balete, D. S., B. R. Tabaranza, Jr., and L. R.
Heaney. 2006. An annotated checklist of the
land birds of Camiguin Island, Philippines, pp.
58-72. In Heaney, L. R., ed., The Mammals and
Birds of Camiguin Island, Philippines, a Distinctive
Center of Biodiversity. Fieldiana Zoology, n s 106:
58-72.

Collar, N. J. 1997. Family Psittacidae (Parrots),

pp. 280-477. In del Hoyo, J., A. Elliott and J.

Sargatal, eds., Handbook of the Birds of the World,

Vol. 4. Lynx Edicions. Barcelona.
Collar. N. J., N. A. D. Mallari, and B. R

Tabaranza. Jr. 1999. Threatened birds of the

Philippines. Bookmark, Makati City.

56

FIELDIANA: ZOOLOGY

Dickinson, E. C, R. S. Kennedy, and K. C. Parkes.
1991. The Birds of the Philippines: An Annotated
Check-List. British Ornithologists' Union Check-
List No. 12. Tring, UK.

Fairbanks, R. G. 1989. A 17,000 glacio-eustatic sea
level record: Influence of glacial melting rates on the
Younger Dryas event and deep-ocean circulation.
Nature, 342: 637-642.

Forshaw, J. M. 1989. Parrots of the World. Lans-
downe Editions, Melbourne.

Funk, D. J., and K. E. Omland. 2003. Species-level
paraphyly and polyphyly: Frequency, causes and
consequences with insights from animal mitochon-
drial DNA. Annual Review of Ecology and System-
atics, 34: 397^423.

Hanebuth, T., K. Stattegger, and P. M. Grootes.
2000. Rapid flooding of the Sunda Shelf: A late-
glacial sea-level record. Nature, 288: 1033-1035.

Heaney, L. R. 1986. Biogeography of mammals in
Southeast Asia: Estimates of rates of colonization,
extinction, and speciation. Biological Journal of the
Linnean Society, 28: 127-165.

— . 1991a. An analysis of patterns of distribution
and species richness among Philippine fruit bats
(Pteropodidae). Bulletin of the American Museum
of Natural History, 206: 145-167.

-. 1991b. A synopsis of climatic and vegetational

change in Southeast Asia. Climatic Change, 19: 53-61 .

Heaney, L. R., and J. C. Regalado, Jr. 1998.
Vanishing Treasures of the Philippine Rain Forest.
The Field Museum, Chicago.

Heaney, L. R., and B. R. Tabaranza, Jr. 1997. A
preliminary report on mammalian diversity and
conservation status of Camiguin Island, Philippines.
Sylvatrop (1995), 5: 57-64.

. 2006a. Introduction to the mammal and land

bird faunas of Camiguin Island, Philippines, pp. 1-
13. In Heaney, L. R., ed., The Mammals and Birds
of Camiguin Island, Philippines, a Distinctive Cen-
ter of Biodiversity, Fieldiana Zoology, n.s., 106:
1-72.

-. 2006b. A new species of forest mouse, genus

Apomys (Rodentia, Muridae) from Camiguin
Island, Philippines, pp. 14-27. In Heaney, L. R.,
ed., The Mammals and Birds of Camiguin Island,
Philippines, a Distinctive Center of Biodiversity.
Fieldiana Zoology, new series, 106: 1-72.
Heaney, L. R., B. R. Tabaranza, Jr., D. S. Balete,
and N. Rigertas. 2006. Synopsis and biogeography
of the mammals of Camiguin Island, Philippines,

pp. 28^8. In Heaney, L. R., ed.. The Mammals and
Birds of Camiguin Island, Philippines, a Distinctive
Center of Biodiversity, Fieldiana Zoology, n.s., 106:
1-72.

IUCN Species Survival Commission. 1994. IUCN Red
list categories, as approved by the 40th meeting of
the IUCN Council, Gland, Switzerland, 30 Novem-
ber 1994. IUCN-The World Conservation Union,
Gland, Switzerland.

Juniper, T., and M. Parr. 1998. Parrots: A Guide to
Parrots of the World. Yale University Press, New
Haven, Connecticut.

Kennedy, R. S., P. C. Gonzales, E. C. Dickinson, H.
C. Miranda, Jr., and T. H. Fisher. 2000. A Guide
to the Birds of the Philippines. Oxford University
Press, Oxford.

Mallari, N. A. D., B. R. Tabaranza, Jr., and M. J.
Crosby. 2001. Key Conservation Sites in the
Philippines. Bookmark, Makati City.

Peterson, A. T. 1996. Geographic variation in sexual
dichromatism in birds. Bulletin of the British
Ornithologists' Club, 116: 156-172.

Peterson, A. T., L. S. Ball, and K. W. Brady. 2000.
Distribution of the birds of the Philippines: Bio-
geography and conservation priorities. Bird Conser-
vation International, 10: 149-167.

Pimentel, R. A. 1979. Morphometries: The Multivar-
iate Analysis of Biological Data. Kendall/Hunt,
Dubuque, Iowa.

Rand, A. L, and D. S. Rabor. 1960. Birds of the
Philippine islands: Siquijor, Mount Malindang,
Bohol, and Samar. Fieldiana Zoology, 35: 221 —
441.

. 1969. New birds from Camiguin South,

Philippines. Fieldiana Zoology, 51: 157-168.

Rickart, E. A., L. R. Heaney, and B. R. Tabaranza,
Jr. 2002. Review of Bull im us (Muridae: Murinae)
and description of a new species from Camiguin
Island, Philippines. Journal of Mammalogy, 83:
421^136.

Smithe, F. B. 1975. Naturalist's Color Guide. The
American Museum of Natural History, New York.

Statsoft Inc. 1995. Statistica for Windows, Statsoft
Inc., Tulsa.

Steppan, S., C. Zawadski, and L. R. Heaney. 2003. A
molecular assessment of phylogenetic relationships
and patterns of phylogenesis in the Philippine murid
rodent Apomys. Biological Journal of the Linnean
Society, 80:699-715.

TELLO ET AL.: A NEW SPECIES OF HANGING-PARROT

57

An Annotated Checklist of the Birds of Camiguin
Island, Philippines

Danilo S. Balete,1'3 Bias R. Tabaranza, Jr.,2 and Lawrence R. Heaney3

Abstract

Fifty-five species of resident breeding and two species of migratory land birds have been
documented on Camiguin Island, including one bittern, one eagle, one junglefowl, two rails,
eight doves and pigeons, one parrot, three cuckoos, one owl, three swiftlets, one dollarbird,
two kingfishers, one bee-eater, one hornbill, one pitta, one triller, two bulbuls, one crow, four
thrushes, three warblers, six flycatchers, one pipit, one wood swallow, two starlings, three
sunbirds, two flowerpeckers, two white-eyes, and two munias. At least seven species reported
here are first records for Camiguin. Ten species are widespread Philippine endemics, two are
near-endemics, and one (Loriculus camiguinensis, described in this volume) is endemic to
Camiguin. Additionally, four endemic subspecies are recognized from Camiguin, Ixos everetti
catarmanensis, Hypothymis azurea catarmanensis, Diceum trigonostigma isidroi, and Zosterops
nigrorum catarmanensis. While this list is not comprehensive, the presence of 57 species
demonstrates that many species were able to cross a narrow but permanent sea channel, and
the presence of four endemic subspecies and one endemic species indicates that some genetic
differentiation has resulted.

Introduction

Although the avifaunas of many of the
Philippine islands have been reported in com-
prehensive fashion (summarized by Dickinson et
al., 1991), no such report has been made for
Camiguin Island, with the result that Camiguin
has not been included in most analyses of avian
biogeography and conservation priorities (e.g.,
Hauge et al., 1986; Dickinson et al., 1991; Collar

1 Laksambuhay Conservation Inc., 10241 Mt. Bu-
lusan Street, U-2. Los Banos, Laguna, Philippines.

2 Department of Biology, Uigan Institute of Tech-
nology, Mindanao State University, Iligan City,
Lanao del Norte, Philippines.

3 Field Museum of Natural History, 1400 South
Lake Shore Drive, Chicago. IL 60605-2496, U.S.A.

et al., 1999; Peterson et al., 2000; but see Mallari
et al., 2001, Ong et al., 2002). We note that
Camiguin Norte, which lies north of Luzon, is
often confused with Camiguin. As documented
in the following list, the land birds of Camiguin
Island include a diverse assemblage of at least 57
species. These records are based on voucher
specimens that were obtained principally during
the 1960s and 1990s and are deposited at the
Delaware Museum of Natural History (dmnh),
the Field Museum of Natural History (fmnh),
and Mindanao State University-Iligan Institute
of Technology (msu-iit), as detailed below and in
Chapter 1 (Heaney & Tabaranza, 2006a). Addi-
tional records based on specimens at the
Museum of Comparative Zoology (mcz), Cam-
bridge, as cited in Dickinson et al. (1991), are
included as well.

58

FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, PP. 58-72

The presence of 55 species of breeding land
birds on an island the size of Camiguin in the
Philippines is not remarkable; indeed, as docu-
mented below, surveys of birds on Camiguin
were not exhaustive, and the current total is well
below the total number likely to be present.
However, given that Camiguin is an oceanic
island surrounded by deep water but only about
10 km from the shore of Mindanao, these data
are a useful indicator of the ability of these 55
species to readily cross a sea channel of this
breadth, given an extensive period of time.
Further, as noted below, the avifauna of
Camiguin is now considered to include four
endemic subspecies of birds (Ixos everetti catar-
manensis, Hypothymis azurea catarmanensis,
Diceum trigono stigma isidroi, and Zosterops
nigrorum catarmanensis) and an endemic species
of parrot {Loriculus camiguinensis; Tello et al.,
2006). This observation indicates that genetic
differentiation has taken place in at least some of
these species, leading in one case to significant
change. It is our hope that this paper will
encourage more extensive and focused analysis
of the avifauna of Camiguin, on both ecology
and evolution.

For notes on methods of collecting and for
a list and description of the collecting sites, see
Heaney and Tabaranza (2006a). As noted in that
chapter, elevations from the 1960s should be
viewed as rough estimates. Species whose names
are listed in brackets are of uncertain documen-
tation and are not included in our tally of species
numbers. Unverified sight records are listed at
the end of this paper. We define "first records"
as those not reported by Dickinson et al. (1991)
and/or Kennedy et al. (2000).

Accounts of Species

Family Ardeidae — Bitterns, Egrets, and Herons

Ixobrychus cinnamomeus — Cinnamon Bittern

The Cinnamon Bittern ranges from India to
China, Ryukyus, Taiwan, and Southeast Asia; in
the Philippines it occurs on most islands, except
the Babuyan and Batanes group, in marshes, rice
fields, and wetlands (Dickinson et al., 1991;
Kennedy et al., 2000). A single specimen was
taken in a highly disturbed agricultural area in
Balbagon, Mambajao, on 28 May 1992, at 10 m,
the first record of this species from the island
(Site 1).

Specimens Examined — Total 1. Site 1 (1
fmnh).

Family Accipitridae — Hawks and Eagles

Spilornis cheela — Crested Serpent-Eagle

The Crested Serpent-Eagle occurs from India
to China, Taiwan, Ryukyus, and Southeast Asia;
it is found throughout the Philippines, except the
Babuyan and Batanes groups of islands, in forest
and forest edge (Dickinson et al., 1991; Kennedy
et al., 2000). Two male specimens were taken on
13 June 1968 on Mt. Catarman, one at 2000 ft
(ca. 600 m; Site 9), the other at an unknown
elevation. We often saw them at nearly all
elevations during fieldwork in 1994 and 1995,
especially near forest.

Specimens Examined — Total 2. Site 9 (1 dmnh;
1 fmnh).

Family Megapodidae — Megapodes and Scrubfowl

[Megapodius cumingii — Tabon]

The Tabon ranges from Sulawesi and Borneo
to the Philippines; once widespread throughout
the country, it is now found mainly in coastal
scrub but also occurs in lowland to montane
forest (Dickinson et al., 1991; Kennedy et al.,
2000). In an earlier enumeration of Philippine
birds, Camiguin was not identified as part of the
Tabon's range (Dickinson et al., 1991). However,
in the more recent compilation, Kennedy et al.
(2000) listed Camiguin as part of the range of
this species without further reference to the
source of their data. Thus, we have not included
M. cumingii in our tally of birds from Camiguin
but note its potential presence.

Family Phasianidae — Partridges, Pheasants,
and Quail

Gallus gallus — Red Junglefowl

The Red Junglefowl occurs from India to
southern China and Southeast Asia, including
virtually all of the Philippines, except the
Batanes group of islands, in forest and forest
edge up to 2000 m (Dickinson et al., 1991;
Kennedy et al., 2000). A male specimen was
taken on Mt. Timpoong at 3150 ft (ca. 950 m)
on 14 June 1969 (Site 13). In May 1994, a male
specimen was taken in a Victor trap in Kital-is,
Sagay at 1200-1400 m (Site 6). We also heard
them in March 1995 crowing close to our
campsite on Mt. Timpoong at 1275 m (Site 7).

Specimens Examined — Total 2. Site 6 (1 msu-
iit); Site 13 (1 dmnh).

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 59

Family Rallidae — Coots, Crakes, Rails,

and Waterhens

Porzana fusca- Ruddy-breasted Crake

The Ruddy-breasted Crake ranges from India
to Japan, Ryukyus, and Southeast Asia; in the
Philippines it has been recorded only on the
islands of Bohol, Cagayancillo, Leyte, Luzon,
Mindanao, Mindoro, Negros, Panay, Samar,
and Sibuyan, usually in marshes and rice fields
but also along forest paths up to 1500 m
(Dickinson et al., 1991; Kennedy et al., 2000).
A male specimen was taken in Kasangsangan in
the vicinity of Mt. Catarman at 1000 ft (ca.
300 m) on 22 June 1968 (Site 11). This is the first
record of this species on Camiguin.

Specimens Examined — Total 1. Site 11 (1

FMNH).

Rallina eurizonoides — Slaty-legged Crake

The Slaty-legged Crake occurs from India and
Sri Lanka to Taiwan, Ryukyus, and Southeast
Asia. In the Philippines it is rather uncommon on
most islands but absent on the Palawan and Sulu
groups of islands, in scrub and primary to
secondary forest up to 900 m (Dickinson et al.,
1991; Kennedy et al., 2000). The record on
Camiguin, a first for this island, consists of two
specimens, both taken in Victor traps, from
Kital-is, Sagay, at 900-1100 m on 19 May 1994
(Site 4) and at 1200-1400 m on 26 May 1994
(Site 6). The following year, another was taken
on Mt. Timpoong in primary montane forest at
1275 m on 23 March 1995 (Site 7).

Specimens Examined — Total 3. Site 4 (1 msu-
iit); Site 6 (1 msu-iit); Site 7 (1 fmnh).

Family Columbidae — Doves and Pigeons
Treron vernans — Pink-necked Green-Pigeon

The Pink-necked Green-Pigeon is widespread
in Southeast Asia; it has been recorded through
most of the Philippines, except the Babuyan and
Batanes groups of islands, principally in lowland
second-growth forest below 300 m (Dickinson,
1991; Kennedy et al., 2000). An adult male
specimen was taken on 29 June 1969 in Puntod,
Mahinog, at 800 ft (ca. 250 m).

Specimens Examined — Total 1. Site 14 (1
dmnii).

Phapitreron leucotis — White-eared Brown-Dove
The White-eared Brown-Dove is endemic to
the Philippines; it occurs throughout the country
with the exception of the Babuyan, Batanes, and
Palawan groups of islands, in primary and

secondary forest up to about 1600 m (Dickinson
et al., 1991; Kennedy et al., 2000). It was
recorded from Mt. Catarman and the nearby
area of Kasangsangan, at 1 000-4500 ft (ca. 300-
1400 m) in June 1968 (Sites 9 and 11). Further
records were obtained from Mt. Timpoong at
3150-4000 ft (ca. 950-1200 m) and the nearby
area of Puntod at 800 ft (ca. 250 m) in June 1969
(Sites 13 and 14, respectively). We also recorded
it in May 1994 in Kital-is, Sagay, at 900-1100 m
(Site 4).

Specimens Examined — Total 12. Site 4 (1 msu-
iit); Site 9 (3 dmnh, 1 fmnh); Site 1 1 (3 fmnh);
Site 13 (2 fmnh); Site 14 (2 fmnh).

Ptilinopus occipitalis — Yellow-breasted Fruit-
Dove

The Yellow-breasted Fruit-Dove is endemic to
the Philippines; it is fairly widespread through-
out the country, except the Babuyan, Batanes,
Palawan, and Sulu groups of islands, in forest up
to 1800 m (Dickinson et al, 1991; Kennedy et
al., 2000). It was recorded on Mt. Catarman and
the nearby areas of Gidag-on and Kasangsangan
at 1000^1500 ft (ca. 300-1400 m) in June 1968
(Sites 9-11). Additional records came from Mt.
Timpoong at 3150^4800 ft (ca. 950-1450 m) in
June 1969 (Sites 12 and 13).

Specimens Examined — Total 18. Site 9 (3
dmnh, 3 fmnh); Site 10 (1 dmnh, 1 fmnh); Site
1 1 (1 fmnh); Site 12 (1 fmnh); Site 13 (4 dmnh, 4
fmnh).

Columba vitiensis — Metallic Pigeon

The Metallic Pigeon occurs on Borneo,
Sulawesi, Moluccas, Lesser Sunda Islands, New
Guinea, and Samoa; it is an uncommon resident
of most islands throughout the Philippines, in
lowland to mossy forest (Dickinson et al., 1991;
Kennedy et al., 2000). Records from Camiguin
were all taken in 1968. A female specimen was
taken from Mt. Catarman at 2000 ft (ca. 600 m)
on 14 June (Site 9) and three adult males and an
adult female from Kasangsangan at 1000-
2000 ft (ca. 300-600 m) in the vicinity of Mt.
Catarman on 11-18 June (Site 11).

Specimens Examined — Total 5. Site 9 (1
fmnh); Site 11 (3 dmnh, 1 fmnh).

Macropygia phasianella — Reddish Cuckoo-Dove
The Reddish Cuckoo-Dove ranges from Bor-
neo, the Moluccas, Sulawesi, and the Lesser
Sunda islands to New Guinea and Samoa; it
occurs throughout the Philippines, in lowland to
mossy forest above 2000 m (Dickinson et al.,

60

FIELDIANA: ZOOLOGY

1991; Kennedy et al., 2000). It was recorded on
Mt. Catarman and the nearby area of Kasang-
sangan at 1000-2500 ft (ca. 300-750 m) in June
1968 (Sites 9 and 11). Additional records came
from Mt. Timpoong at 3150-5700 ft (ca. 950-
1700 m) and the nearby area of Puntod at 800 ft
(ca. 250) in June 1969 (Sites 12-14). In May
1994, we recorded it in Kital-is, Sagay, at 1000-
1400 m (Sites 5 and 6).

Specimens Examined — Total 30. Site 5 (1 msu-
iit); Site 6 (2 msu-iit); Site 9 (1 fmnh); Site 1 1 (3
dmnh, 3 fmnh); Site 12 (9 dmnh, 7 fmnh); Site 13
(1 dmnh, 2 fmnh); Site 14 (1 fmnh).

Streptopelia bitorquata — Island Collared-Dove

The Island Collared-Dove is found from
Borneo and the Lesser Sunda islands to Java;
in the Philippines it is a fairly uncommon
resident of most islands, except the Batanes
group of islands, mainly in relatively open fields
in the lowlands and sometimes in mangrove
(Dickinson et al., 1991; Kennedy et al., 2000).
Two male specimens were collected from Pun-
tod, Mahinog, at 700-800 ft (ca. 200-250 m),
one of which was a juvenile taken at 800 ft (ca.
250 m) on 28 June 1968 (Site 14).

Specimens Examined — Total 2. Site 14 (1
dmnh, 1 fmnh).

Geopelia striata — Zebra Dove

The Zebra Dove ranges from Southeast Asia
to Australia; it occurs throughout the Philip-
pines, except the Babuyan and Batanes groups of
islands, in open country and cultivated areas in
the lowlands (Dickinson et al., 1991; Kennedy et
al., 2000). One of us (B.R.T.) netted one some
50 m away from the beach in Manuyog, Sagay,
on 10 May 1994 (Site 3), but it was released. No
other specimen was recorded of this species in
1994. It was not recorded during our fieldwork in
1992 and 1995.

Specimens Examined — None.

Chalcophaps indica — Common Emerald-Dove

The Common Emerald-Dove is found from
India and Sri Lanka to China, Taiwan, Ryukyus
and Southeast Asia, New Guinea, and Australia;
it occurs throughout the Philippines, often in
second-growth forest in the lowlands up to
1000 m (Dickinson et al., 1991; Kennedy et al.,
2000). Records on Camiguin consist of speci-
mens taken in Kasangsangan, near Mt. Catar-
man, at 1000-2500 ft (ca. 300-750 m) in June
1968 (Site 11) and from Puntod at 800 ft (ca.
250 m) in May 1969 (Site 14).

Specimens Examined — Total 5. Site 11 (2
dmnh, 2 fmnh); Site 14 (1 fmnh).

Family Psittacidae — Lorikeets, Cockatoos, Parrots,
and Racquet-tails

Loriculus camiguinensis — Camiguin Hanging-
Parrot

Described as a new species in this volume
(Tello et al., 2006), this is the only species of bird
currently recognized as endemic to Camiguin. Its
elevational range, based on specimens collected
in the 1960s, is 300 to 1350, thus including
lowland and montane rain forest. Most of this
habitat has been destroyed on Camiguin, leading
to concern for its conservation status.

Specimens Examined — Total 23. Site 9 (2
dmnh); Site 11 (5 dmnh, 5 fmnh); Site 13 (5
dmnh, 6 fmnh).

Family Cuculidae — Cuckoos, Malkohas, and Coucals

Cacomantis variolosus — Brush Cuckoo

The Brush Cuckoo occurs from Southeast
Asia to Australia and the Southwest Pacific; it is
found throughout the Philippines, except the
Babuyan and Batanes groups of islands, from
mangrove to mossy forest at 2000 m (Dickinson
et al., 1991; Kennedy et al., 2000). The records
from Camiguin were taken from Mt. Catarman
at 2500 ft (ca. 750 m) in June 1968 (Site 9) and
from Mt. Timpoong at 3150^1800 ft (ca. 950-
1450 m) in June 1969 (Site 13). We also recorded
it in Kital-is, Sagay, at 900-1100 (Site 4) and
1200-1400 m in May 1994 (Site 6).

Specimens Examined — Total 7. Site 6 (1 msu-
iit); Site 9 (1 fmnh); Site 13 (3 dmnh, 2 fmnh).

Eudynamys scolopacea — Common Koel

The Common Koel occurs from India through
Southeast Asia to northern Australia; it is found
throughout the Philippines, in primary and
secondary lowland forest (Dickinson et al.,
1991; Kennedy et al., 2000). The single specimen
from Camiguin was taken on Mt. Timpoong at
3150 ft (ca. 950 m) on 17 June 1969 (Site 13).

Specimens Examined — Total 1. Site 13 (1
fmnh).

Centropus viridis — Philippine Coucal

The Philippine Coucal is endemic to the
Philippines, where it occurs throughout the
country, except the Palawan group of islands,
from grassland to forest at 2000 m (Dickinson et
al., 1991; Kennedy et al., 2000). The records on
Camiguin consist of specimens taken from

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 61

Kasangsangan in the vicinity of Mt. Catarman at
1000 1500 it (ca. 300 450 m) in June 1968 (Site
11) and from Mt. Timpoong at 3150 ft (ca.
950 m) in June 1969 (Site 13).

Specimens Examined— Total 6. Site 11 (1
dmnii. 2 fmnh); Site 13(1 dmnh, 2 fmnh).

Family Strigidae — Owls

Ninox philippensis Philippine Hawk-Owl

The Philippine Hawk-Owl is endemic to the
Philippines, occurring throughout most of the
islands, except the Babuyan. Batanes, and
Palawan groups, in the lowlands up to 1 800 m
(Dickinson et al., 1991; Kennedy et al., 2000).
The records from Camiguin consist of a female
taken from Mt. Catarman at 1500 ft (ca. 450 m)
on 17 June 1968 (Site 9) and another female from
Mt. Timpoong on 13 June 1969 at an unknown
elevation. Specimens of other species from Mt.
Timpoong on the same date were taken at
3150 ft (ca. 950 m).

Specimens Examined — Total 2. Site 9 (1
fmnh); Site 13(1 fmnh).

Family Apodidae — Swifts

Collocalia mearnsi — Philippine Swiftlet

The Philippine Swiftlet is endemic to the
Philippines, where it is recorded mainly on the
islands of Bohol, Cebu, Luzon, Mindanao,
Mindoro, Negros, Palawan, and Panay, usually
in forest and forest clearings above 900 m; it is
absent from the Babuyan, Batanes, and Sulu
groups of islands (Dickinson et al., 1991;
Kennedy et al., 2000). Specimens from Camiguin
were taken from Mt. Catarman and in the
nearby Kasangsangan at 1000-2000 ft (ca. 300-
600 m) on 16 June 1968 (Sites 9 and 11) and
from Mt. Timpoong on 23 June 1969 from an
unknown elevation. Specimens of other species
from this site were taken at 3100-3350 ft (ca.
950-1000 m), with several at 4000 ft (ca.
1200 m).

Specimens Examined — Total 4. Site 9 (1
fmnh); Site 11 (1 fmnh); Site 13 (2 fmnh).

Collocalia esculenta — Glossy Swiftlet

The Glossy Swiftlet occurs from the Anda-
mans, Nicobars. and Malay Peninsula to New
Guinea and the southwest Pacific; it is found
throughout the Philippines, from sea level to
mountaintops (Dickinson et al., 1991; Kennedy
et al., 2000). It was commonly recorded on Mt.
Catarman and in the nearby area of Kasangsan-

gan at 1000-2000 ft (ca. 300-600 m) in June
1968 (Sites 9 and 11) and on Mt. Timpoong at
3150-4800 ft (ca. 950-1450 m) in June 1969
(Sites 12 and 13).

Specimens Examined — Total 31. Site 9 (1
fmnh); Site 11 (2 dmnh, 7 fmnh); Site 12 (2
dmnh, 1 fmnh); Site 13 (9 dmnh, 9 fmnh).

Collocalia troglodytes— Pygmy Swiftlet

The Pygmy Swiftlet is endemic to the Philip-
pines, where it occurs throughout most of the
country, except the Babuyan, Batanes, and Sulu
groups of islands, usually at low to middle
elevations in forested areas (Dickinson et al.,
1991; Kennedy et al., 2000). The records from
Camiguin were taken on 13 and 21 June 1968 in
Kasansangan at 1500-2000 ft (ca. 450-600 m)
and on Mt. Timpoong at 3150 ft (ca. 950 m) on
25 June 1969 (Site 13).

Specimens Examined — Total 4. Site 11 (3
dmnh); Site 13 (1 fmnh).

Family Coraciidae — Rollers

Eurystomus orientalis — Dollarbird

The Dollarbird occurs from India to New
Guinea and the southwest Pacific, including all
of the Philippines, except the Batanes group of
islands, in forest edge and clearings in the
lowlands up to 1200 m (Dickinson et al., 1991;
Kennedy et al., 2000). Dickinson et al. (1991)
noted a single specimen deposited in the Museum
of Comparative Zoology, Harvard University,
taken from Mambajao on 19 August 1921; no
elevation was indicated.

Specimens Examined — None.

Family Alcedinidae — Kingfishers

Ceyx lepidus — Variable Dwarf-Kingfisher

The Variable Dwarf-Kingfisher occurs from
the Moluccas to New Guinea and the southwest
Pacific islands; in the Philippines it is recorded
on the islands in the Central Philippines as well
as on the Mindanao and Sulu groups of islands,
in primary and secondary lowland forest (Dick-
inson et al, 1991; Kennedy et al., 2000). It was
recorded from Mt. Timpoong at 3150 ft (ca.
950 m) on 17 and 18 June 1969 (Site 13). In May
1994, two individuals were recorded in Kital-is,
Sagay, at 900-1 100 m (Site 4), including one that
was taken in a Victor trap on 17 May.

Specimens Examined — Total 4. Site 4 (2 msu-
iit); Site 13 (1 dmnh, 1 fmnh).

62

FIELDIANA: ZOOLOGY

Halcyon Moris — White-collared Kingfisher

The White-collared Kingfisher occurs widely
from northeast Africa to southern China, Ryu-
kyus, Southeast Asia to New Guinea, Australia,
and the southwest Pacific; it is found nearly all
over the Philippines, from exposed reefs to open
country and forest edge (Dickinson et al., 1991;
Kennedy et al., 2000). It was recorded in Gidag-
on and Kasangsangan, both in the vicinity of Mt.
Catarman, at 500-2500 ft (ca. 150-600 m; Sites
10 and 1 1) in June 1968 and on Mt. Timpoong at
3150 ft (ca. 950 m; Site 13), in Puntod at 800 ft
(ca. 250 m; Site 14), and Mantigue Island (Site
19) in June 1969. We recorded it further in
Balbagon, Mambajao, at 10 m in May 1992 (Site
1). On 10 May 1994, we also recorded it in
Manuyog, Sagay, at 80 m (Site 3).

Specimens Examined — Total 33. Site 1 (1
fmnh); Site 10 (6 dmnh, 4 fmnh); Site 11 (1
dmnh, 1 fmnh); Site 13 (5 dmnh, 5 fmnh); Site 14
(4 dmnh, 5 fmnh); Site 19 (1 dmnh).

Family Meropidae — Bee-Eaters

Merops viridis — Blue-throated Bee-eater

The Blue-throated Bee-eater occurs in South-
east Asia and the Philippines, where it is known
from many islands, except the Babuyan, Batanes,
Palawan, and Sulu groups; it often found in open
country, scrubs, and forest clearings (Dickinson
et al., 1991; Kennedy et al., 2000). On Camiguin,
it was recorded from Mt. Catarman and in
nearby Kasangsangan at 1000-4500 m (ca. 300-
1400 m) in June 1968 (Sites 9 and 1 1) and on Mt.
Timpoong at 3350 ft (ca. 1000 m) in June 1969
(Site 13).

Specimens Examined — Total 9. Site 9 (1 dmnh,
2 fmnh); Site 11 (2 dmnh, 2 fmnh); Site 13 (1
dmnh, 1 fmnh).

Family Bucerotidae — Hornbills

Aceros leucocephalus — Writhed Hornbill

The Writhed Hornbill is endemic to Mind-
anao, Dinagat, and Camiguin, in forest up to
1200 m (Dickinson et al., 1991; Kennedy et al.,
2000). The records from Camiguin in 1968
consist of a male taken from Kasangsangan in
the vicinity of Mt. Catarman at 1000 ft (ca.
300 m) on June 16 (Site 11) and another male
from Mt. Catarman at 2000-4000 ft (ca. 600-
1200 m) on June 17 (Site 9). A female was also
obtained from Mt. Timpoong at 3150 ft (ca.
950 m) on 16 June 1969 (Site 13). We did not
record it during our fieldwork in the 1990s.
Mallari et al. (2001) mention, without further

reference to the source, a juvenile captured in
Ginsiliban in 1993. The most recent evaluation
of this species found it to be fairly common
despite "losing ground clearly to habitat clear-
ance, hunting and trapping for trade" and
assigned it a Near-Threatened status (Collar et
al., 1999).

Specimens Examined — Total 3. Site 9 (1
dmnh); Site 11 (1 fmnh); Site 13 (1 fmnh).

Family Pittidae — Pittas

Pitta erythrogaster — Red-bellied Pitta

The Red-bellied Pitta occurs from Sulawesi to
the Moluccas and New Guinea and northeast
Australia; it is found throughout the Philippines,
except the Batanes group of islands, in primary
and secondary forest usually below 1000 m
(Dickinson et al., 1991; Kennedy et al., 2000).
Three individuals, a male and two females, were
taken from Mt. Timpoong at 3150-5700 ft (ca.
950-1700 m) on 14-25 June 1969 (Sites 12 and
13). We recorded it further on the same
mountain at 1475 m on 22 March 1995 (Site 8).
In May 1994, it was the most common bird taken
in Victor traps in Kital-is, Sagay, at 900-1 100 m
to 1200-1400 m (Sites 4-6).

Specimens Examined — Total 9. Site 4 (2 msu-
iit); Site 5 (1 msu-iit); Site 6 (2 msu-iit); Site 8 (1
fmnh); Site 12 (1 dmnh); Site 13 (2 fmnh).

Family Campephagidae — Cuckoo-Shrikes, Minivets,
and Trillers

Lalage nigra — Pied Triller

The Pied Triller ranges from the Nicobars to
Southeast Asia; it occurs throughout the Philip-
pines, except the Batanes and Babuyan groups of
islands, usually in open areas in the lowlands up
to 1400 m (Dickinson et al., 1991; Kennedy et
al., 2000). It was recorded from Gidag-on and
Kasangsangan in the vicinity of Mt. Catarman at
500-2000 ft (ca. 150-600 m) in June 1968 (Sites
10 and 1 1). In June of the following year, records
came mainly from Mt. Timpoong at 3150 ft (ca.
950 m; Site 13), Puntod, at 800 ft (ca. 150 m; Site
14), and Mantigue Island (Site 19).

Specimens Examined — Total 30. Site 10 (9
dmnh, 4 fmnh); Site 11 (2 fmnh); Site 13 (1
dmnh); Site 14 (4 dmnh, 5 fmnh); Site 19 (5
dmnh).

Family Pycnonotidae — Bulbuls

Pycnonotus goiavier — Yellow-vented Bulbul

The Yellow-vented Bulbul occurs widely in
Southeast Asia and throughout the Philippines,

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 63

except the Babuyan and Batanes groups of
islands, in scrub and second-growth forest
(Dickinson et al., 1991: Kennedy et al., 2000).
It was recorded from Mt. Catarman and its
nearby areas of Gidag-on and Kasangsangan at
500-4000 ft (ca. 150 1200 m) in June 1968 (Sites
9-11). In June 1969, additional records were
obtained from Mt. Timpoong at 3150-4800 ft
(ca. 950-1450 m; Sites 12 and 13) and the nearby
area of Puntod at 800 ft (ca. 250 m; Site 14) as
well as from Mantigue Island on 28 June (Site
19). We recorded it further in Balbagon,
Mambajao. at 10 m on 28 May 1992 (Site 1).
Dickinson et al. (1991) noted the presence of
another specimen from an unspecified locality on
Camiguin, deposited at the Museum of Com-
parative Zoology, Harvard University.

Specimens Examined — Total 30. Site 1 (1
fmnh); Site 9 (1 dmnh, 1 fmnh); Site 10 (5 dmnh,
5 fmnh); Site 11 (2 dmnh, 3 fmnh); Site 12 (1
dmnh); Site 13 (5 fmnh); Site 14 (2 dmnh, 3
fmnh); Site 19 (1 dmnh).

Ixos everetti — Yellowish Bulbul

The Yellowish Bulbul is endemic to the islands
of Biliran, Bucas Grande, Dinagat, Leyte, Mind-
anao, Panaon, Samar, and Siargao, the Sulu
group, and Camiguin, usually in primary and
secondary lowland forest (Dickinson et al., 1991;
Kennedy et al., 2000). Three subspecies are
recognized, including /. e. catarmanensis, which
is restricted to Camiguin (Rand & Rabor, 1969).
Our comparison of specimens at fmnh with those
of other named subspecies supports the distinc-
tiveness of the Camiguin population; it is sub-
stantially the largest and darkest among the three
races that are currently recognized (Dickinson et
al.. 1991; Kennedy et al., 2000). The call of this
species is equally distinct (Kennedy et al., 2000).
The geographic variation observed in this species
as currently defined is unusually great and is
suggestive of a potential species group. Further
taxonomic studies are warranted.

This species was recorded in June 1968 from
Mt. Catarman and the nearby areas of Gidag-on
and Kasangsangan at 500-4500 ft (ca. ISO-
MOO m; Sites 9-11). Additional records were
obtained in June 1969 from Mt. Timpoong at
3150-5700 ft (ca. 950-1700 m; Sites 12 and 13)
and the nearby area of Puntod at 800 ft (ca.
250 m; Site 14). In May 1994, it was one of the
most common birds mist netted in Kital-is,
Sagay, at 900-1 100 m to 1200-1400 m (Sites 4-
6).

Specimens Examined— Total 94. Site 4 (8 msu-
iit); Site 5 (2 msu-iit); Site 6 (1 msu-iit); Site 9 (6
dmnh, 15 fmnh); Site 10 (6 dmnh); Site 11 (9
dmnh, 7 fmnh); Site 12 (2 dmnh, 2 fmnh); Site 13
(20 dmnh, 13 fmnh); Site 14 (2 dmnh, 1 fmnh).

Family Corvidae — Crows

Corvus /w/oY>/7nvn/?o.v— Large-Billed Crow

The Large-Billed Crow occurs from Iran to
Northeast Asia, China, Taiwan, Ryukyus, and
Southeast Asia; it is found nearly throughout the
Philippines, in forest edge to plantations and in
towns (Dickinson et al., 1991; Kennedy et al.,
2000). A male specimen was taken from Gidag-
on in the vicinity of Mt. Catarman at 1000 ft (ca.
300 m) on 24 June 1968 (Site 10).

Specimens Examined — Total 1. Site 10 (1
fmnh).

Family Turdidae — Robins, Shamas, and Thrushes
Copsychus saularis — Oriental Magpie-Robin

The Oriental Magpie-Robin occurs from
Pakistan and India to southern China and
Southeast Asia; in the Philippines, it occurs
throughout the country, except the Palawan,
Babuyan, and Batanes groups of islands, usually
in second-growth and scrubby forest (Dickinson
et al., 1991; Kennedy et al., 2000). It was
recorded from Mt. Catarman and in the nearby
area of Gidag-on at 500 3000 ft (ca. 150-900 m)
in June 1968 (Sites 9 and 10). Additional records
were obtained in June 1969 from Mt. Timpoong
at 4000 ft (1200 m; Site 13) and the nearby area
of Puntod at 800 ft (ca. 250 m; Site 14). We
recorded it further in May 1994 in Kital-is,
Sagay, at 900-1 100 m (Site 4) and 1200-1400 m
(Site 6).

Specimens Examined — Total 16. Site 4 (3 msu-
iit); Site 6 (1 msu-iit); Site 9 (1 fmnh); Site 10 (4
dmnh, 3 fmnh); Site 13 (3 fmnh); Site 14 (1
fmnh).

Saxicola caprata — Pied Bushchat

The Pied Bushchat occurs from Iran to
southwest China, Southeast Asia, and New
Guinea; it is a common resident in the Philip-
pines, except the Sulu, Palawan, Babuyan, and
Batanes groups of islands, usually in fairly dry,
open country (Dickinson et al., 1991; Kennedy et
al., 2000). A female specimen was taken from
Mt. Catarman at 2000 ft (ca. 600 m) on 19 June
1968 (Site 9).

Specimens Examined — Total 1. Site 9 (1
fmnh).

64

FIELDIANA: ZOOLOGY

Zoothera andromedae — Sunda Ground-Thrush

The Sunda Ground-Thrush occurs from Java
and Sumatra to the Lesser Sunda Islands; it is an
uncommon resident in the Philippines, where it
has been recorded mainly on mountains in the
northern and central Luzon, Mindanao, Mind-
oro, Negros, and Panay, in the understory of
forest above 1000 m (Dickinson et al., 1991;
Kennedy et al., 2000). We recorded a single
specimen, taken in a Victor rat trap, from Mt.
Timpoong at 1275 m on 23 March 1995 (Site 7);
this is the first record for Camiguin.

Specimens Examined — Total 1. Site 7 (1
fmnh).

Zoothera dauma — Scaly Ground-Thrush

The Scaly Ground-Thrush is an uncommon
winter visitor to the Philippines from Siberia,
India, China, Japan, Korea, and Taiwan. It has
been recorded previously only on the islands of
Batan, Catanduanes, Fuga, Luzon, Marinduque,
Mindoro, Palawan, and Sibuyan, where it
forages on the ground and forest understory at
all elevations (Dickinson et al., 1991; Kennedy et
al., 2000). Our record on Camiguin consists of
a single specimen taken in a Victor rat trap on
Mt. Timpoong at 1275 m on 19 March 1995 (Site
7). This is the first record of this species for the
island.

Specimens Examined — Total 1. Site 7 (1
fmnh).

Family Sylviidae — Old World Warblers

Phylloscopus trivirgatus — Mountain Leaf-War-
bler

The Mountain Leaf- Warbler occurs from the
Malay Peninsula, Borneo, Sumatra, Java, and
the Lesser Sunda Islands to New Guinea; in the
Philippines it is found mainly on the islands of
Luzon, Mindanao, Negros, Palawan, and Panay,
in montane and mossy forest above 800 m
(Dickinson et al., 1991; Kennedy et al., 2000).
An endemic subspecies, P. t. diuatae, is recog-
nized on Camiguin and Mt. Hilong-hilong
(eastern Mindanao; Dickinson et al., 1991;
Kennedy et al., 2000), though Dickinson (in
litt.) considered this in need of reassessment.
Records on Camiguin were obtained from Mt.
Catarman at 2000^950 ft (ca. 600-1500 m) in
June 1968 (Site 9) and from Mt. Timpoong at
4800-5700 ft (ca. 1450-1700 m) in June 1969
(Sites 12 and 13). It was also recorded in Kital-is,
Sagay, at 900-1100 m in May 1994 (Site 4).

Specimens Examined— Total 31. Site 4 (1 msu-
iit); Site 9 (3 dmnh, 5 fmnh); Site 12 (10 dmnh,
10 fmnh); Site 13 (2 dmnh).

Megalurus timoriensis — Tawny Grassbird

The Tawny Grassbird occurs from the Lesser
Sunda Islands, Moluccas, and Sulawesi to New
Guinea and Australia; it is found throughout
most of the Philippines, excluding the Babuyan,
Batanes, Palawan, and Sulu groups of islands, in
grasslands and disturbed forest up to 2000 m
(Dickinson et al., 1991; Kennedy et al., 2000). It
was recorded in 1968 from Kasangsangan in the
vicinity of Mt. Catarman at 1000 ft (ca. 300 m)
on June 20 (Site 11). Additional records were
obtained from Mt. Timpoong at 3250 ft (ca.
980 m) in June 1969 (Site 13). We also recorded
it in May 1994 in Kital-is, Sagay, at 1000-
1300 m (Site 5).

Specimens Examined — Total 10. Site 5 (1 msu-
iit); Site 11 (2 fmnh); Site 13 (3 dmnh, 4 fmnh).

Cisticola exilis — Bright-capped Cisticola

The Bright-capped Cisticola ranges from India
to southern China, Taiwan, Southeast Asia
(excluding the Malay Peninsula) to New Guinea,
Australia, and the southwest Pacific; it occurs on
most islands of the Philippines, excluding the
Babuyan, Batanes, Palawan, and Sulu groups of
islands, in grassy habitats and rice fields (Dick-
inson et al., 1991; Kennedy et al., 2000). Two
adult males were taken in 1968, one from Mt.
Catarman at 2000 ft (ca. 600 m) on 19 June (Site
9) and the other from Kasangsangan in the
vicinity of Mt. Catarman at 1500 ft (ca. 450 m)
on 21 June (Site 11).

Specimens Examined — Total 2. Site 9 (1
dmnh); Site 11 (1 fmnh).

Family Muscicapidae — Flycatchers

Ficedula hyperythra — Snowy-browed Flycatcher
The Snowy-browed Flycatcher ranges from
the Himalayas to southern China, Taiwan, and
Southeast Asia; it occurs on most of the larger
islands throughout the Philippines, in dense
forest understory usually above 1000 m, but is
absent on Sibuyan and the Batanes and Sulu
groups of islands (Dickinson et al., 1991;
Kennedy et al., 2000). On Camiguin, we
recorded this species for the first time in Kital-
is, Sagay, at 900-1100 m in May 1994 (Site 4)
and on Mt. Timpoong at 1275 m on 25 March
1995 (Site 7).

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 65

Specimens Examined— Total 5. Site 4 (4 nisi
in); Site 7 (1 fmnh).

Ficedula westermanni Little Pied Flycatcher

The Little Pied Flycatcher occurs from India
to southern China and Southeast Asia; in the
Philippines, it has been recorded only from
Luzon, Mindanao, Mindoro, Negros, southern
Palawan, and Panay, in forest and forest edge
above 800 m. usually in middle story and forest
canopy (Dickinson et al., 1991; Kennedy et al.,
2000). The records from Camiguin were taken
from Mt. Catarman at 2000^950 ft (ca. 600-
1500 m) in June 1968 (Site 9) and on Mt.
Timpoong at 3150-5700 ft (ca. 950-1700 m) in
June 1969 (Sites 12 and 13). On 29 May 1992, we
recorded it on Mt. Mambajao at 1000 m (Site 2)
and in May 1994 in Kital-is, Sagay, at 900-
1300 m (Sites 4 and 5).

Specimens Examined — Total 42. Site 2 (1
fmnh); Site 4 (3 msu-iit); Site 5 (1 msu-iit); Site
9(12 dmnh, 10 fmnh); Site 12 (9 dmnh, 4 fmnh);
Site 13 (2 fmnh).

Cyornis rufigastra — Mangrove Blue Flycatcher

The Mangrove Blue Flycatcher ranges from
the Malay Peninsula, Java, Sumatra, and Sula-
wesi to Borneo and the Philippines; it occurs
throughout most of the islands of the Philip-
pines, except the Batanes group, usually in
disturbed forest habitat (Dickinson et al., 1991;
Kennedy et al., 2000). It was recorded from Mt.
Catarman and the nearby areas of Gidag-on and
Kasangsangan at 500^1950 ft (ca. 150-1500 m)
in June 1968 (Sites 9-11). Further records were
obtained from Mt. Timpoong at 3150-4800 ft
(ca. 950-1450 m; Sites 12 and 13) and the nearby
area of Puntod at 800 ft (ca. 250 m; Site 14) in
June 1969. We also recorded it in Kital-is, Sagay,
at 900-1 100 m in May 1994 (Site 4).

Specimens Examined — Total 70. Site 4 (4 msu-
iit); Site 9 (8 dmnh, 9 fmnh); Site 10 (7 dmnh, 4
fmnh); Site 11 (6 dmnh, 6 fmnh); Site 12 (2
dmnh, 4 fmnh); Site 13 (7 dmnh, 5 fmnh); Site 14
(4 dmnh, 4 fmnh).

Rhipidura javanica — Pied Fantail

The Pied Fantail ranges from the Malay
Peninsula, Sumatra, and Java to Borneo and
the Philippines, where it occurs throughout most
of the islands, except the Babuyan and Batanes
groups, usually in lowland second growth,
residential areas, bamboo thickets, and man-
groves (Dickinson et al., 1991; Kennedy et al.,

2000). It was recorded mainly from the lower
slopes of Mt. Catarman and the nearby areas of
Gidag-on and Kasangsangan in June 1968 at
500-2000 ft (ca. 150-600 m; Sites 9 11). Addi-
tional records in June 1969 were taken from Mt.
Timpoong, at 3150 ft (ca. 950 m; Site 13) and the
nearby area of Puntod at 800 ft (ca. 250 m; Site
14). We also recorded it in May 1994 in Kital-is,
Sagay, at 1000-1400 m (Sites 5 and 6).

Specimens Examined — Total 32. Site 5 (2 msu-
iit); Site 6 (1 msuiit); Site 9 (2 fmnh); Site 10 (7
dmnh, 7 fmnh); Site 1 1 (3 dmnh, 3 fmnh); Site 13
(1 dmnh, 2 fmnh); Site 14 (2 dmnh, 2 fmnh).

Terpsiphone cinnamomea — Rufous Paradise-Fly-
catcher

The Rufous Paradise-Flycatcher occurs
throughout the Philippines, excluding the Babu-
yan, Batanes, and Palawan groups of islands, in
understory of forest up to 1200 m; it also occurs
on several of the Talaud Islands, Indonesia
(Dickinson et al., 1991; Kennedy et al., 2000).
Three specimens, two females and a male, were
taken on 16-23 June 1969 from Mt. Timpoong at
1500-3150 ft (ca. 450-950 m; Site 13); another
female was recorded from Puntod at 800 ft (ca.
250 m) on 27 June 1969 (Site 14). In May 1994, it
was recorded in Kital-is, Sagay, at 900-1100 m
(Site 4).

Specimens Examined — Total 5. Site 4 (1 msu-
iit); Site 13 (1 dmnh, 2 fmnh); Site 14 (1 dmnh).

Hypothymis azurea — Black-naped Monarch

The Black-naped Monarch ranges from India
to southern China and Taiwan to Southeast
Asia; it occurs nearly throughout the Philippines,
except the Batanes group of islands, usually in
disturbed forest and forest edge below 1500 m
(Dickinson et al., 1991; Kennedy et al., 2000).
The population on Camiguin is recognized as
a distinct endemic subspecies, H. a. catarmanen-
sis (Rand & Rabor, 1969). Records from
Camiguin in June 1968 came from Mt. Catarman
and the nearby areas of Gidag-on and Kasang-
sangan at 1000^1950 ft (ca. 300-1450 m). Fur-
ther records in June 1969 came from both Mt.
Catarman at 2000-4950 ft (ca. 600-1450 m; Sites
9-11) and Mt. Timpoong at 3150 ft (ca. 950 m;
Site 13). In May 1994, we recorded it in Kital-is,
Sagay, at 900-1 100 m to 1200-1400 m (Sites 4-
6).

Specimens Examined — Total 33. Site 4 (2 msu-
iit). Site 5 (1 msu-iit); Site 6 (2 msu-iit); Site 9 (3

66

FIELDIANA: ZOOLOGY

dmnh, 4 fmnh); Site 10 (1 dmnh, 1 fmnh); Site 1 1
(2 dmnh, 3 fmnh); Site 13 (12 dmnh, 2 fmnh).

Family Motacillidae — Pipits, Wagtails

Anthus hodgsoni — Olive Tree-Pipit

The Olive Tree-Pipit is an uncommon migrant
from eastern Asia to India, southern China,
Taiwan, Ryukyus, and Southeast Asia; in the
Philippines it has been recorded on many of the
larger islands, except the Batanes and Sulu
groups, usually in pine and oak forest above
300 m (Dickinson et al., 1991; Kennedy et al.,
2000). On Camiguin, we recorded this species in
primary montane forest on Mt. Timpoong at
1275 m on 24 March 1995 (Site 7); it is the first
record from Camiguin.

Specimens Examined — Total 1. Site 7 (1
fmnh).

Family Artamidae — Wood Swallows

Artamus leucorynchus — White-breasted Wood-
Swallow

The White-breasted Wood-Swallow occurs
from Borneo, Sulawesi, to New Guinea, Aus-
tralia, and the southwest Pacific; it occurs
throughout the Philippines, except the Babuyan
and Batanes groups of islands, usually at forest
edge or in disturbed forest up to 1800 m
(Dickinson et al., 1991; Kennedy et al., 2000).
Records from Camiguin in June 1968 came from
Mt. Catarman and the nearby areas of Gidag-on
and Kasangsangan at 500-2000 ft (ca. ISO-
GOO m; Sites 9 and 10). Additional records in
June 1969 were from Mt. Timpoong at 3150 ft
(ca. 950 m; Site 13) and Puntod at 800 ft (ca.
250 m; Site 14).

Specimens Examined — Total 19. Site 9 (1
dmnh, 1 fmnh); Site 10 (5 dmnh; 4 fmnh); Site
11 (1 dmnh); Site 13 (2 dmnh; 4 fmnh); Site 14 (1
dmnh).

Family Sturnidae — Mynas and Starlings

Aplonis panayensis — Asian Glossy Starling

The Asian Glossy Starling is found from
eastern India to Southeast Asia; it occurs
throughout the Philippines, in agricultural and
residential areas and forest edge in the lowlands
(Dickinson et al., 1991; Kennedy et al., 2000).
Records from Camiguin in June 1968 came from
the lower slopes of Mt. Catarman and nearby
areas of Gidag-on and Kasangsangan at 500-
2000 ft (ca. 150-600 m; Sites 10 and 11). Two
other specimens were obtained in Puntod at
800 ft. (ca. 250 m) in June 1969 (Site 14). We

recorded it further in Balbagon, Mambajao, on
28 May 1992 at 10 m (Site 1).

Specimens Examined — Total 40. Site 1 (1
fmnh); Site 10 (15 dmnh, 21 fmnh); Site 11 (1
dmnh); Site 14 (2 dmnh).

Sarcops calvus — Coleto

The Coleto occurs almost exclusively in the
Philippines, in agricultural plantations, second
growth, and forest edge in the lowlands; its only
record outside the country is on Banggi Island,
off Borneo (Dickinson et al., 1991; Kennedy et
al., 2000). Within the Philippines itself, however,
it is absent from the Babuyan, Batanes, and
Palawan groups of islands (Dickinson et al.,
1991; Kennedy et al., 2000). In June 1968, two
specimens each were taken from Gidag-on at
500-1500 ft (ca. 1 50^450 m; Site 10) and Ka-
sangsangan at 1000 ft (ca. 300 m; Site 11), both
in the vicinity of Mt. Catarman.

Specimens Examined — Total 4. Site 10 (2
dmnh); Site 1 1 (2 fmnh).

Family Nectariniidae — Spiderhunters and Sunbirds

Anthreptes malacensis — Plain-throated Sunbird

The Plain-throated Sunbird occurs throughout
most of Southeast Asia; in the Philippines, it is
found on almost all islands, except the Babuyan
and Batanes groups, in coconut groves, man-
groves, and second-growth forest in the lowlands
(Dickinson et al., 1991; Kennedy et al., 2000).
Records on Camiguin came mainly from Ka-
sangsangan in the vicinity of Catarman Moun-
tain at 1500-2000 ft (ca. 450-600 m) in June
1968 (Site 1 1) and Puntod at 800 ft (ca. 250 m) in
June 1969 (Site 14). A single specimen was taken
on Mt. Timpoong at an unknown elevation on
16 June 1969. Specimens of other species from
the same site and date were taken at 1500-
3150 ft (ca. 450-950 m).

Specimens Examined — Total 12. Site 11 (3
dmnh, 3 fmnh); Site 13 (1 dmnh); Site 14 (5
dmnh).

Nectarinia jugularis — Olive-backed Sunbird

The Olive-backed Sunbird occurs from South-
east Asia to New Guinea, Australia, and the
southwest Pacific; in the Philippines, it occurs on
almost all islands, except the Babuyan and
Batanes groups, usually in heavily disturbed
habitats, including towns and cities, below
1000 m (Dickinson et al., 1991; Kennedy et al.,
2000). Records from Camiguin were obtained
from Mt. Catarman and the nearby areas of

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 67

Gidag-on and Kasangsangan at 500 3000 ft (ca.
150-900 m) in June 1968 (Sites 10 and 11). In
June 1969, additional records came from Mt.
Timpoong at 1000-5000 ft (ca. 300-1500 m;
Sites 12 and 13) and the nearby area of Puntod
at 800 ft (ca. 250 m; Site 14) as well as from the
island of Mantigue (Site 19). We recorded it
further in Balbagon, Mambajao, on 28 May
1992 at 10 m (Site 1) and in Kital-is, Sagay, at
1000-1300 m in May 1994 (Site 5).

Specimens Examined — Total 94. Site 1 (2
fmnh); Site 5 (1 msu-iit); Site 10 (24 dmnh, 20
fmnh); Site 11 (4 dmnh, 6 fmnh); Site 12 (5
dmnh, 3 fmnh); Site 13 (1 fmnh); Site 14 (14
dmnh, 6 fmnh); Site 19 (8 dmnh).

Nectarinia sperata — Purple-throated Sunbird

The Purple-throated Sunbird occurs from
India and Southeast Asia; it is found throughout
the Philippines, except the Batanes group of
islands, in cultivated areas, mangroves, and
secondary forest in the lowlands (Dickinson et
al., 1991; Kennedy et al., 2000). Records from
Camiguin in June 1968 were obtained from Mt.
Catarman and the nearby areas of Gidag-on and
Kasangsangan at 500-3000 ft (ca. 150-900 m;
Sites 9-11). In June the following year, records
were obtained from Mt. Timpoong at 3150 ft
(ca. 950 m; Site 13) and in the nearby area of
Puntod at 800 ft (ca. 250 m; Site 14). Addition-
ally, Dickinson et al. (1991) noted two specimens
from an unspecified locality and elevation on
Camiguin that are deposited at the Museum of
Comparative Zoology, Harvard University.

Specimens Examined — Total 39. Site 9 (3
dmnh, 3 fmnh); Site 10 (6 dmnh, 6 fmnh); Site
11 (3 dmnh, 7 fmnh); Site 13 (7 fmnh); Site 14 (1
dmnh, 3 fmnh).

Family Dicaeidae — Flowerpeckers
Dicaeum trigonostigma — Orange-bellied Flower-
pecker

The Orange-bellied Flowerpecker ranges east-
ern India to Southeast Asia; it occurs throughout
the Philippines, except on the Babuyan and
Batanes groups of islands, in forest edge, second
growth, and cultivated areas below 1500 m
(Dickinson et al., 1991; Kennedy et al., 2000).
On Camiguin, this species is represented by an
endemic subspecies, D. t. isidroi (Rand & Rabor,
1969). Records from Camiguin were obtained
from Mt. Catarman and the nearby areas of
Gidag-on and Kasangsangan at 500-4500 ft (ca.
150-1400 m) in June 1968 (Sites 9-11). Addi-

tional records in June 1969 were obtained from
Mt. Timpoong at 800-3150 ft (ca. 250-950 m:
Site 13) and the nearby area of Puntod at 800 it
(ca. 250 m; Site 14). We obtained a further
record of it in Balbagon, Mambajao, at 10 m on
28 May 1992 (Site 1) and in Kital-is, Sagay, at
900-1 100 m and 1200-1400 m in May 1994
(Sites 4 and 6, respectively).

Specimens Examined — Total 63. Site 1 (1
fmnh); Site 4 (6 msu-iit); Site 6 (5 msu-iit); Site
9 (5 dmnh, 4 fmnh); Site 10 (3 dmnh, 4 fmnh);
Site 11 (5 dmnh, 11 fmnh); Site 13 (8 dmnh, 3
fmnh); Site 14 (6 dmnh, 2 fmnh).

Dicaeum pygmaeum — Pygmy Flowerpecker

The Pygmy Flowerpecker is common through-
out the Philippines, except Mindoro and the
Babuyan, Batanes, and Sulu groups of islands, in
forest, forest edge, and second growth mainly
below 1000 m (Dickinson et al., 1991; Kennedy
et al., 2000). A single record exists from
Camiguin, taken on 13 June 1968 in Kasangsan-
gan, in the vicinity of Mt. Catarman at an
unknown elevation. A specimen of Columba
vitiensis was obtained on the same date and
from the same site at 1000 ft (ca. 300 m).

Specimens Examined — Total 1. Site 11 (1
dmnh).

Family Zosteropidae — White-eyes

Zosterops everetti — Everett's White-eye

Everett's White-eye ranges from the Malay
Peninsula, Borneo, and Talaud to the Philip-
pines; it occurs only on the islands southeast of
Luzon, from Samar to Mindanao and Cebu to
the Sulu and Tawi-tawi groups of islands, in
scrub and forest up to 1000 m (Dickinson et al.,
1991; Kennedy et al., 2000). Records from
Camiguin were obtained from Mt. Catarman
and the nearby areas of Gidag-on and Kasang-
sangan at 500^1950 ft (ca. 150-1500 m) in June
1968 (Sites 9 11). Further records were obtained
in June 1969 from Mt. Timpoong at 3150 ft (ca.
950 m; Site 13) and the nearby area of Puntod at
800 ft (ca. 250 m; Site 14).

Specimens Examined — Total 52. Site 9 (4
dmnh, 1 fmnh); Site 10 (11 dmnh, 9 fmnh); Site
11 (4 dmnh, 9 fmnh); Site 13 (4 dmnh, 8 fmnh);
Site 14 (2 fmnh).

Zosterops nigrorum — Yellowish White-eye

The Yellowish White-eye is endemic to the
Philippines, where it occurs on Luzon, Mindoro.
Negros, and Panay and the adjacent small

FIELDIANA: ZOOLOGY

islands, in forest and forest edge below 1000 m; it
is absent from the Babuyan, Mindanao, Pala-
wan, and Sulu groups of islands (Dickinson et
al., 1991; Kennedy et al., 2000). The population
on Camiguin is the southernmost extension of
this species' range and is treated as an endemic
subspecies, Z. n. catarmanensis (Rand & Rabor,
1969). Records from Camiguin in June 1968 were
all obtained from Mt. Catarman at 2000^1950 ft
(ca. 600-1500 m; Site 9). Additional records in
June 1969 were all obtained from Mt. Timpoong
at 3150-5400 ft (ca. 950-1600 m; Sites 12 and
13). We recorded it further on Mt. Mambajao at
1000 m on 29 May 1992 (Site 2) and in Kital-is,
Sagay, at 900-1 100 m and 1200-1400 m in May
1994 (Sites 4 and 6, respectively).

Specimens Examined — Total 181. Site 2 (2
fmnh); Site 4 (4 msu-iit); Site 6 (3 msu-iit); Site 9
(53 dmnh, 39 fmnh); Site 12 (36 dmnh, 15 fmnh);
Site 13 (20 dmnh, 9 fmnh).

Family Estrildidae — Avadavat, Parrotfinches,
and Munias

Lonchura leucogastra — White-Bellied Munia

The White-Bellied Munia ranges from the
Malay Peninsula, Sumatra, Borneo, and
throughout the Philippines, in forest, forest edge,
grassland, and in rice farms (Dickinson et al.,
1991; Kennedy et al., 2000). Records of this
species on Camiguin were all obtained in Gidag-
on and Kasangsangan, near Mt. Catarman, at
500-1000 ft (ca. 150-300 m) in June 1968 (Sites
10 and 11).

Specimens Examined — Total 6. Site 10 (1
dmnh, 3 fmnh); Site 11 (1 dmnh, 1 fmnh).

Lonchura malacca — Chestnut Munia

The Chestnut Munia is recorded from India
and Nepal to southwestern China, Taiwan, and
Southeast Asia and is widespread throughout the
Philippines, associated mainly with rice fields,
grasslands, and open country. The only records
from Camiguin were from Kasangsangan near
Mt. Catarman at 1500 ft. (ca. 450 m) on 21-22
June 1968 (Site 11).

Specimens Examined — Total 3. Site 11 (1
dmnh, 2 fmnh).

Sight Records

Following the recommendation appearing in
the Rules for New Records (Kennedy et al.,
2000), we have deliberately omitted from the
Species Accounts all sight records that were not

verified or supported by two other observers
after the two-year rule for publication. The
following is the list of such species, including
species identified by calls only, which when
verified would constitute additional records for
Camiguin and is indicative of the potentially far
richer avifauna of Camiguin than the current
Species Accounts would suggest. We hope that
this listing will encourage other ornithologists
and bird-watchers to conduct more studies of
Camiguin birds.

Family Accipitridae

Accipiter sp.

Observed by one of us (B.R.T.) along a trail
leading to Site 4 on 12 March 1994 between 6 am
and 3:30 pm.

Family Columbidae

Ducula poliocephala — Pink-necked Pigeon

Observed by one of us (B.R.T.) along the same
trail where an Accipiter was sighted (see above),
on the same date and time.

Family Cuculidae

Cacomantis merulinus — Plaintive Cuckoo

The call of this cuckoo was noted by B.R.T. at
Site 4 on 13 May 1994 as one of the most
commonly heard birdcalls at this site. The local
name of the bird is pitokai.

Family Strigidae

Otus sp.

The call of an unidentified scops-owl was
noted by B.R.T. at Site 4 on 14 May 1994 at ca.
5:40 am.

Family Oriolidae

Oriolus sp.

The call of an unidentified oriole was noted by
B.R.T. at Site 4 on the same date and time that
the scops-owl was heard (see above).

Family Pachycephalidae

Pachycephala sp.

The call of an unidentified whistler was noted
by B.R.T. at Site 4 on the same date and time
that the scops-owl was heard (see above).

Discussion

The bird fauna of Camiguin is currently
represented by 57 species, consisting of 55

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 69

resident breeders and two migrants in 26 bird
families. Seven species are reported here for the
first time: Ixobrychus cinnamomeus, Porzana
fusca, Rallina eurizonoides, Zoothera androme-
dae, Zoothera dauma, Ficedula hyperythra, and
Anthus hodgsoni. Doves and pigeons (Columbi-
dae) are the most diverse family on Camiguin,
with eight species recorded on the island,
followed by flycatchers (Musicapidae) with six
species, and thrushes (Turdidae) with four
species. The remaining 22 bird families are
represented by one to three species only. Because
Camiguin is a volcanic island of recent geological
origin and is separated from mainland Mind-
anao by a sea channel that is only 10 km wide
but 385 m deep, it would have not been
connected to Mindanao during the many periods
of low sea level in the Pleistocene (Heaney, 1986;
Heaney & Tabaranza, 2006a). Thus, the current
species assemblage reached Camiguin Island by
colonization, and their presence on Camiguin is
documentation of the capacity of these species
for overwater dispersal.

A brief comparison of the birds on Camiguin
with those on the closest islands, and sources of
potential colonizers, including Mindanao and
the central Philippine islands of Negros, Panay,
and Cebu, is revealing. Except for the endemic
Camiguin Colasisi, or Hanging-Parrot, Loriculus
camiguinensis, 53 of the remaining species of
resident birds are also found on Mindanao;
Zosterops nigrorum does not occur on Mind-
anao. With Negros, Panay, and Cebu, Camiguin
shares 50, 47, and 42 of its resident bird species,
respectively (Dickinson et al., 1991; Kennedy et
al., 2000). Luzon, which is farthest away from
Camiguin in terms of potential source of
colonizers, also shares with it at least 50 resident
bird species (Dickinson et al., 1991; Kennedy et
al., 2000). These data indicate a remarkably high
faunal similarity with Mindanao, though it does
not provide clear evidence of biogeographic
affinity, given an equally high degree of similar-
ity with Luzon and Panay, for instance. Thus, it
appears that Camiguin has been colonized by
widespread species.

The presence of at least one species endemic to
Camiguin {Loriculus camiguinensis, described in
this volume) and four endemic subspecies of
birds clearly indicates that colonization rates
have been so low, even with its proximity to
Mindanao, that substantial genetic differentia-
tion has occurred. In this regard, it is noteworthy
that none of the other islands adjacent to

Mindanao, such as Basilan, Bohol, Dinagat.
Leyte, Samar, or Siargao, regardless of area and
distance from it, has a single-island endemic
(Dickinson et al., 1991; Kennedy et al.. 2000;
Peterson et al., 2000). All these islands formed
part of the Pleistocene island of Greater Mind-
anao and thus were repeatedly connected by
dryland areas during much of the Pleistocene
(Heaney, 1986, 2000; Heaney & Regalado, 1998;
Steppan et al., 2003), unlike Camiguin, which
remained isolated (Heaney & Tabaranza, 2006a).

The current listing of birds on Camiguin is far
from comprehensive, pending more systematic
field surveys, but based on the quite extensive
collection effort during the late 1960s, it is
worthwhile to note here the apparent absence
or depauperateness of several families that are
otherwise well represented on other oceanic
islands in the Philippines such as Sibuyan
(currently, the most comprehensively studied
small oceanic island in the Philippines for birds;
see Goodman et al., 1995). Sibuyan also provides
a good contrast with Camiguin in terms of the
number of species that have successfully colo-
nized oceanic islands in the Philippines. Good-
man et al. (1995) identified at least 102 resident
species on Sibuyan; Camiguin's current list pales
in comparison. Among raptors, for instance,
only one eagle, S. cheela (Accipitridae), and one
owl, N. philippensis (Strigidae), have been
recorded on Camiguin. Sibuyan, in contrast,
has at least six species of eagles, three species of
owls, and one species of falcon (Falconidae)
present. A similar trend is apparent in rails
(Rallidae), swiftlets (Apodidae), and kingfishers
(Alcedinidae). Also noteworthy is the poor
representation of the larger doves (Columbidae)
on Camiguin that occur on Sibuyan, such as
Ducula poliocephala, D. carola, D. aena, and
larger parrots (Psittacidae), such as Priorniturus
discurus and Tanygnatlnts lucionensis (Goodman
et al., 1995).

On the other hand, Camiguin and Sibuyan
appear to share the absence of species that are
generally associated with wetlands and freshwa-
ter swamps (Anhingidae, Threskiornithidae, and
Jacanidae) as well as species in high-elevation
forest habitats, such as Orthotomus cucullatus,
Bradypterus caudatus, Rhynomyias goodfellowi,
Eumyias panayensis, Ficedula crypta, Culicicapa
helianthea, Serinus estherae, and Pyrrhula leuco-
genis, which occur on some of the larger islands,
including Mindanao (Dickinson et al., 1991;
Goodman et al., 1995; Kennedy et al., 2000).

70

FIELDIANA: ZOOLOGY

Both islands also lack some general forest and
forest-edge resident birds in the following
families: Podargidae, Trogonidae, Capitonidae,
Chloropseidae, Paridae, and Sittidae and Lanii-
dae, Dicruridae, Oriolidae, Rhabdornithidae,
and Timaliidae. Camiguin also currently lacks
records of Turnicidae, Rostratulidae, Scolopaci-
dae, Tytonidae, Caprimulgidae, Hemiprocnidae,
Picidae, Eurylaimidae, Hirundinidae, Alaudidae,
Oriolidae, and Pachycephalidae.

It is further interesting to note that on
Camiguin, in contrast to Sibuyan (Goodman &
Gonzales, 1990; Goodman et al., 1995) and other
larger and more speciose islands, several wide-
spread species associated with open country,
cultivated areas, and disturbed habitats in the
lowlands, such as Pycnonotus goiavier, Rhipidura
javanica, and Nectarinia jugularis, are commonly
found in relatively intact montane and mossy
forest up to the peaks of Mt. Timpoong. A
similar pattern in mammals was earlier observed
on Negros, which is a relatively depauperate
island in terms of mammal diversity, where
several commensal species {Suncus murinus,
Rattus exulans, and R. tanezumi) are present
and often abundant at all elevations and in all
habitat types (Heideman et al., 1987; Heaney et
al., 1989), and on Camiguin, where Suncus mur-
inus is abundant in montane forest (Heaney &
Tabaranza, 1997; Heaney et al., 2006). The docu-
mentation of the above pattern of distribution
along elevational gradients has led to the hypo-
thesis that the number of species in the native
community of small mammals in mature forest
determines the success of non-native small mam-
mals on oceanic islands (Heaney et al., 1999).
Might this apply to birds on Camiguin as well? Or
might habitat disturbance be an equally important
factor contributing to this trend in bird diversity
and distribution on Camiguin? Camiguin is
a volcanically active island, with recorded erup-
tions of at least two of its volcanos (Hibok-hibok
and Vulcan) within recent times. Mt. Hibok-hibok
erupted in 1827, 1862, 1871-1875, 1897, 1902,
and 1948-1958 and Vulcan in 1871 and 1874
(http://www.volcano.si.edu/gvp/world/volcano.
cfm?vnum= 070 1-08; http://hannover.park.org/
Philippines/pinatubo/page9.html). This suggests
that the island has had bouts of habitat
disturbance brought about by these eruptions,
aside from the ongoing human-induced habitat
alteration and fragmentation (Heaney & Tabar-
anza, 2005a). Further surveys of the avifauna
of Camiguin will enable a more comprehensive

analysis of the factors that influence diversity
and distribution patterns of birds among
oceanic islands in the Philippines.

Finally, we note that Camiguin is the home
not only to a newly described parrot (Loriculus
camiguinensis; Tello et al., 2006) and four
endemic races of birds (Ixos everetti catarma-
nensis, Hypothymis azurea catarmanensis, Di-
ceum trigonostigma isidroi, and Zosterops ni-
grorum catarmanensis) but also to at least two
species of mammals (Heaney & Tabaranza,
2006b), one amphibian, and seven plants (Hea-
ney et al., 2006). It clearly warrants protection as
a unique portion of the natural heritage of
the Philippines. We strongly recommend active
protection of the remaining forest. This will
benefit the wildlife of the island but will also be
essential to the economic and social stability of
Camiguin because the mountain forests are the
source of the island's essential and precious
water. This and other conservation issues are
discussed in more detail in Heaney and Tabar-
anza (2006a).

Acknowledgments

For assistance with fieldwork on Camiguin,
we thank N. Antoque, E. Batara, N. Batocael,
N. Bojo, M. Carmona, A. DeOcampo, R.
Fernandez, M. Jayoma, L. Mostrales, A. T.
Peterson, G. Rosell, A. Tabaranza, B. Tabaranza
III, and D. Tabaranza. Permits were provided by
the Protected Areas and Wildlife Bureau (Phi-
lippine Department of Environment and Natural
Resources); we especially thank A. C. Alcala, C.
Custodio, M. Mendoza, and W. Pollisco. We are
indebted to D. Willard for access to specimens at
the Field Museum and especially to G. Hess for
loans of specimens and information on collec-
tions under his care at the Delaware Museum.
We wish to thank Sean Bober and Sarah Lansing
for assistance with preparing the manuscript. We
are grateful to D. Willard and J. Bates for helpful
comments on earlier drafts and to N. J. Collar
and E. C. Dickinson for insightful reviews of the
manuscript. Fieldwork was supported by the
World Environment and Resources Program of
the John D. and Catherine T. MacArthur
Foundation. Additional support was provided
by the Marshall Field Fund, Ellen Thorne Smith
Fund, and Barbara Brown Fund for Mammal
Research of the Field Museum.

\LETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 71

Literature Cited

Collar, N. J., N. A. Mallari, and B. R. Tabaranza.
Jr. 1999. Threatened Birds of the Philippines.
Bookmark, Makati City.

Dickinson, E. C, R. S. Kennedy, and K. C. Parkes.
1991. The birds of the Philippines: An annotated
checklist. British Ornithologists' Union Checklist
No. 12.

Goodman, S. M., D. E. Willard, and P. C. Gonzales.
1995. The birds of Sibuyan Island, Romblon
Province, with particular reference to elevational
distribution and biogeographic affinities. Fieldiana:
Zoology, n.s., 82: 1-57.

Goodman, S. M., and P. C. Gonzales. 1990. The birds
of Mt. Isarog National Park, southern Luzon,
Philippines, with particular reference to altitudinal
distribution. Fieldiana: Zoology, n.s., 60: 1-39.

Hauge, P., J. Terborgh, B. Winter, and J. Parkin-
son. 1986. Conservation priorities in the Philippines.
Forktail, 2: 83-91.

Heaney. L. R. 1986. Biogeography of mammals in
Southeast Asia: Estimates of rates of colonization,
extinction, and speciation. Biological Journal of the
Linnean Society, 28: 127 165.

— . 2000. Dynamic disequilibrium: a long-term,
large-scale perspective on the equilibrium model of
island biogeography. Global Ecology and Biogeog-
raphy. 9: 59-74.

Heaney. L. R.. D. S. Balete, E. A. Rickart. R. C. B.
Utzurrum. and P. C. Gonzales. 1999. Mammalian
diversity on Mount Isarog, a threatened center of
endemism on Southern Luzon Island. Philippines.
Fieldiana: Zoology, n.s., 95: 1-62.

Heaney, L. R., P. D. Heideman, E. A. Rickart, R. B.
Utzurrum, and J. S. H. Klompen. 1989. Elevational
zonation of mammals in the central Philippines.
Journal of Tropical Ecology. 5: 259-280.

Heaney, L. R., and J. C. Regalado, Jr. 1998.
Vanishing Treasures of the Philippine Rain Forest.
The Field Museum. Chicago.

Heaney. L. R., and B. R. Tabaranza, Jr. 1997.
Preliminary report on mammalian diversity and
conservation status in Camiguin Island. Philippines.
Sylvatrop(1995). 5: 57-64.

— . 2006a. Mammal and land bird studies on
Camiguin Island, Philippines: Background and
conservation priorities, pp. 1-13. /// Heaney, L.
R., ed.. The Mammals and Birds of Camiguin
Island, Philippines, a Distinctive Center of Bio-
diversity. Fieldiana Zoology, n.s., 106: 1-72.

— . 2006b. A new species of forest mouse, genus
Apomys (Mammalia. Rodentia) from Camiguin
Island. Philippines, pp. 14-27. In Heaney. L. R..
ed.. The Mammals and Birds of Camiguin Island.
Philippines, a Distinctive Center of Biodiversity.
Fieldiana Zoology, n.s.. 106: 1-72.

Heaney. L. R., B. R. Tabaranza. Jr.. D. S. Balete,
and N. Rigertas. 2006. Synopsis and biogeography
of the mammals of Camiguin Island, Philippines.
pp. 28-48. In Heaney, L. R., ed.. The Mammals and
Birds of Camiguin Island. Philippines, a Distinctive
Center of Biodiversity. Fieldiana Zoology, n.s.. 106:
1-72.

Heideman, P. D., L. R. Heaney. R. L. Thomas, and
K. R. Ericsson. 1987. Patterns of faunal diversity
and species abundance of non-volant mammals on
Negros Island, Philippines. Journal of Mammalogy,
68: 884-888.

Kennedy, R. S.. P. C. Gonzales, E. C. Dickinson, H.
C. Miranda. Jr.. and T. H. Fisher. 2000. Guide to
the Birds of the Philippines. Oxford University
Press, New York.

Mallari, N. A. D, B. R. Tabaranza. Jr.. and M. J.
Crosby. 2001. Key Conservation Sites in the
Philippines. Bookmark, Makati City.

Ong, P., L. E. Afuang.and R. G. Rosell-Ambal. eds.
2002. Philippine Biodiversity Conservation Priori-
ties: A Second Iteration of the National Biodiversity
Strategy and Action Plan. Philippine Department of
the Environment and Natural Resources, Quezon
City.

Peterson. A. T.. L. G. Ball, and K. W. Brady. 2000.
Distribution of birds of the Philippines: biogeogra-
phy and conservation priorities. Bird Conservation
International. 10: 149-167.

Rand, A. L., and D. S. Rabor. 1969. New birds from
Camiguin South. Philippines. Fieldiana Zoology, 51:
157-168.

Steppan, S., C. Zawadski, and L. R. Heaney. 2003. A
molecular assessment of phylogenetic relationships
and patterns of phylogenesis in the Philippine murid
rodent Apomys. Biological Journal of the Linnean
Society, 83: 699-715.

Tello, J. G.. J. F. Degner. J. M. Bates, and D. E.
Willard. 2006. A new species of Hanging-Parrot
(Aves: Psittacidae: Loriculus) from Camiguin
Island. Philippines, pp. 58-72. //; Heaney. L. R.,
ed.. The Mammals and Birds of Camiguin Island.
Philippines, a Distinctive Center of Biodiversity.
Fieldiana Zoology, n.s.. 106: 1-72.

72

FIELDIANA: ZOOLOGY

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