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Full text of "The Mammals and birds of Camiguin Island, Philippines, a distinctive center of biodiversity"

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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 



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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 km 2 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. 9 r 10.5'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, 124 r 45.5'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 — X A 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/km 2 in 1980 to 
306 persons/km" in 2000: these were well above 
the national density of 160 persons/km 2 and 
251 persons/km 2 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. 



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104-166. 

Mitchell, A. H. G.. F. Hernandez, and A. P. Dela 
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MlTTERMEIER, R. A., N. MYERS, P. RoBLES GlL, AND C. 

G. Mittermeier. eds. 1999. Hotspots, Earth's 
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Mittermeier. eds.. 1997. Megadiversity: Earth's 
Biologically Wealthiest Nations. CEMEX. Monter- 
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Mlsser. G. G.. L. R. Heaney. and B. R. Tabaranza, 
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National Mapping and Resource Information Au- 
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Ong. P.. L. E. Afiang. andR. G. Rosell-Ambal, eds. 
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Peterson, A. T.. L. G. Ball, and K. W. Brady. 2000. 
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12 



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Primack, R. B. 1998. Essentials of Conservation 
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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. Heaney 1 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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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 M 1 (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 35 c 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. 



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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 Rigertas 1 



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 km 2 , 

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 km 2 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 





*=1960s 
■=1990s 



11 



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maximum elevation 



J 1 1 1. ■ 1 1 

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II 



0) 



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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 





3 (0.3) 


1 (0.3) 


0* 


1 (0.2) 





5 


Sanctis marinas 


2(1.4) 


0* 


0* 


10(2.9) 


56(8.5) 


9 (2.3) 


77 


Apomys camiguinensis 





14(1.5) 


0* 


2 (0.6) 


9(1.4) 





25 


Bullimus gatnay 





4 (0.4) 


2 (0.6) 


3 (0.9) 


10(1.5) 


1 (0.3) 


20 


C 'runomys melanius 





2 (0.2) 


1 (0.3) 


0* 


2 (0.3) 





5 


Rattas everetti 





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) 





7 


Rattas tanczumi 


21 (14.2) 


8 (0.9) 


4(1.2) 


1 (0.3) 








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 





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 



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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) 





liar prion vcteris whitehead! 


1 (0.17) 











0* 


Macroglossus minimus 


15 (2.5) 


1 (0.33) 


4(0.17) 


9 (0.64) 





Picnot hints jagori 


8(1.33) 





16(0.67) 


1 (0.07) 





Total fruit bats 


51 


5 


35 


14 





Total net-nights 


6 


3 


24 


14 


8 


Fruit bats per net-night 


8.5 


1.67 


1.46 


1.00 






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 



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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. 



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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. Willard 1 



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 



FIELDIANA: ZOOLOGY 



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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 45.2 c /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. 



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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. Heaney 3 



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 



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