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Zoology NEW SERIES, NO. 106

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

Lawrence R. Heaney, Editor

April 5, 2006 Publication 1537

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY UNIVERSITY I LIBR/ AT URBANA-CHA BIOL: -S4*^||

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

Philippine Islands have been recognized This volume contains the results of our studies Thefor over a century as having a fauna that is of Camiguin Island, a small island that lies just characterized by large numbers of endemic and north of central Mindanao. It is easily over- highly distinctive species. For most of this looked on a map of the Philippines, yet, as period, the mammals and birds were thought to shown here, the island supports an endemic be well known, based on studies conducted in the species of small parrot and two endemic species period 1880-1950. However, the description of of small mammals, all discovered in the course of new species during the 1980s implied that our studies. Announcements in the Philippine of the in diversity was higher than had been believed press discovery of the mammals 1994 and suggested that there might be additional and 1995 played an important role in encourag- the declaration of the rain forest localized centers of endemism yet to be found. In ing remaining on as a national a addition, it was during the 1980s that the extent Camiguin park, movement that is to It is our of rain-forest habitat destruction became appar- continuing gain support. hope ent and increased the need for extensive and that this publication will help guide the planning for this area and its when detailed documentation of biological diversity to protected management it has been initiated. The data contained guide development and conservation for the formally here constitute a baseline from which benefit of both wildlife and the human popula- changes be measured in the future and also make tion of the Philippines. may clear how much additional is needed. With this in mind, the authors of this volume study Although Camiguin is a small island, it repre- and their colleagues began investigations of sents an ideal natural laboratory in which to places likely to be of special interest and initiated investigate many aspects of the evolution and a series of publications in Fieldiana: Zoology to conservation of biological diversity. document the findings. The field studies have The assistance of individuals is acknowl- been conducted in a standardized fashion to many edged in each of the chapters that follow, but we allow direct and meaningful comparison between must give special recognition here to the study areas so that geographic patterns of species Philippine Department of Environment and richness between islands and mountain ranges Natural Resources for providing both permits and along elevational gradients could be docu- and encouragement; we especially thank Dr. mented. However, older museum collections, Angel C. Alcala, Dr. Corazon Catibog-Sinha, which often were unreported in the published Mr. Carlo C. Custodio, Atty. Wilfredo Pollisco, literature, have also been utilized, increasing the and Dr. Mundita Lim for their steadfast support. rate of our progress in understanding the re- This project has benefited greatly from the markable mammal and bird faunas of the financial support provided by the John D. and The first in Fieldiana Philippines. publications Catherine T. MacArthur Foundation, from the concerned the mammals of and Leyte nearby Ellen Thorne Smith and Marshall Field Funds of islands north of Mindanao, the birds and the Field Museum, and especially the Barbara mammals of Mt. National Park in Isarog Brown Fund for Mammal Research of the Field southern Luzon, and the birds of Sibuyan Island. Museum. Plans are in place for similar publications on the mammals and birds of the Kitanglad Range of L. R. Heaney northern Mindanao and the mammals of Sibu- November 2005 yan Island, and others are likely to follow. 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 remarkably large number of endemic species

(e.g., Dickinson et al., 1991; Mittermeier et al., The mammal and bird faunas of the Philippine 1997, 1999; Heaney & Regalado, 1998; Heaney et Islands are remarkable for the high total di- al., 1998; Stattersfield et al., 1998). For example, versity of the fauna and especially for the at least 512 of the 898 species of breeding terrestrial vertebrates (57%) are endemic to the Philippines, an unusually high value (Heaney &

1 Most of the endemic to Field Museum of Natural History, 1400 South Regalado, 1998). species Lake Shore Drive, Chicago, IL 60605-2496, U.S.A. the country occur on the large islands of Luzon, 2 Mindanao, Mindoro, Negros, and Palawan (e.g., Department of Biology, Iligan Institute of Tech- et et nology, Mindanao State University, Iligan City, Hauge al., 1986; Heaney, 1986; Heaney al., Lanao del Norte, Philippines. 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 report on the mammals (Heaney, 1984) conclud- restricted to some of the smaller islands as well ed that the island had no endemic mammal

(e.g.. Heaney. 1986: Goodman & Ingle, 1993: species and that it was depauperate relative to its Heaney & Tabaranza. 1997; Musser et al., 1998; area. Subsequent studies on other islands made Heaney & Mallari, 2002: Rickart et al.. 2002). us suspect that those earlier mammal surveys Evidence has suggested that those small islands were incomplete because so few mammal species with endemic species are nearly always islands had been obtained and because the number of surrounded by deep water (more than 125 m). so nonvolant mammal voucher specimens was small that they were not connected to adjacent larger (thus indicating limited sampling effort). To islands during periods of low sea level during the investigate the hypothesis that the previously "ice ages" of the middle and late Pleistocene, measured species richness of mammals was low when sea level dropped to no more than 120 m because of incomplete surveys, we returned to below present level (Heaney. 1986. 1991a.b: Camiguin briefly in 1992 and more extensively in Heaney & Regalado. 1998: Hanebuth et al., 1994 and 1995 to conduct additional mammal 2000; Siddall e^l., 2003). Clearly, if we are to inventories in all the major habitats along the understand the overall patterns of the evolution elevational gradient, especially by trapping small and ecology of diversity in the Philippines, we mammals at higher elevations (Heaney & Taba- must understand the biodiversity patterns of ranza. 1997), and we obtained some new data on these smaller centers of endemism as well as the the birds as well. In this volume, we summarize larger ones. results from both the 1960s and the 1990s One of the smaller Philippine islands sur- surveys and present evidence that both endemic rounded by deep water and still retaining mammal and bird species are present on the moderate rain-forest cover is Camiguin, located island (Heaney et al., 2006; Tello et al., 2006). about 10 km off the north-central shore of There is a second reason that we conducted Mindanao Island, with minimum intervening surveys on Camiguin in the mid-1990s. While the water depth of 385 m (the island of Camiguin Philippines is increasingly being recognized as Norte, which lies north of Luzon, is often a global center for biodiversity, with unusually confused with Camiguin). It is a steeply moun- high levels of endemism. it has simultaneously tainous 2 island (Fig. 1) of about 265 km with vaulted into public view as one of the most several active volcanic cones that reach to severely deforested of the tropical countries and a maximum elevation of about 1620 m. A series home to what may be the greatest concentration of biological surveys on Camiguin in the late of endangered species of mammals and birds 1960s (described below) that focused on birds (Collar et al., 1994, 1999; Heaney & Regalado, also yielded some mammal specimens. An earlier 1998: Stattersfield et al., 1998; Mittermeier et al..

FIELDIANA: ZOOLOGY 124° 38' 124° 40' 124° 42' 124° 44' 124° 46' 124° 48'

9° 15' White Island

9°13

-9°11

Mantigue Island o

9°09

9 07'

9°05'

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), a portion of the country where the 2000; Ong et al., 2002). Satellite maps of forest concentration of endangered mammals and birds cover from 1986 (National Mapping and Re- is especially high (Wildlife Conservation Society source Information Authority, 1988) showed of the Philippines, 1997; Collar et al., 1999; a substantial area of forest cover in the center of Heaney & Mallari, 2002). Because deforestation Camiguin (Fig. 2); such forest cover is now a rarity has generally proceeded rapidly all over the in the Visayas (the islands of the central Philip- country in recent decades, we felt a sense of pines) and adjacent regions (Heaney & Regalado, urgency to obtain current information. Simulta- 1998; Environmental Science for Social Change, neously, the prospect of finding species of

HEANEY AND TABARANZA: MAMMAL AND LAND BIRD STUDIES 124°42'

- 9M5'

ST13'

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 within historic time (Hamilton. 1979; Mitchell et remained to support stable populations of any al., 1986; Hall, 1998, 2002), with the most recent endemic or endangered species was quite exciting, serious eruptions in 1871 and 1951 (Agoo, 1995). posing the potential for positive action and the Five major and several secondary volcanic peaks development of successful conservation pro- (Figs. 1-3) dominate the island's landscape and grams. We therefore proceeded to Camiguin with serve as a tourist attraction; hikers often reach a sense of cautious optimism and found condi- the crater lake in Mt. Hibok-hibok and visit the tions that confirmed all our best hopes but also steam vents along its sides. some of our fears, as we shall describe here. Camiguin is the northernmost, somewhat isolated portion of the Central Mindanao Arc of volcanic activity, which includes areas from Geology, Vegetation, and Climate Mt. Apo in the south to Mts. Kalatungan and Kitanglad in the center and Camiguin in the Camiguin is composed almost entirely of north (Sajona et al., 1997). Volcanic activity in Quaternary volcanic material from currently this arc originated about 2.5 million years ago active volcanoes, and eruptions have occurred (Ma), though most has taken place since about

FIELDIANA: ZOOLOGY 1.25 Ma. Camiguin is probably derived from Rhododendron, Medinilla, and Vaccinium. Gym- a source of that first single magma produced nosperms are conspicuously absent. . . . The undersea lavas in the late Pliocene or early vegetation at the peak is devoid of tall trees Pleistocene (ca. 2 Ma), with dryland appearing and shrubs. It is composed mostly of turfs of not earlier than 1 Ma. Potassium-argon dating grasses." Descriptions of vegetation at our shows the oldest dated magmatic materials on sampling areas during the 1990s are below. the island to be those of the Mt. Ginsiliban and Agoo (1995) noted the presence of the Mt. volcanoes in Butay the southeastern corner following plants endemic to Camiguin: Miquelia of the island which date to 0.34 (Fig. 1), Ma. The reticulata (Icacinaceae), Medinilla multiflora, and volcanic flows peaks associated with Mt. Memecylon subcaudatum (Melastomataceae), Sy- that form Mambajao much of the center of the zigium camiguense (Myrtaceae), Coelogyne con- island are much younger, and Mt. Hibok-hibok fusa, and Goodyera ramosii (Orchidaceae). in the northwestern of the island is the portion A small frog (Oreophryne nana) was the only and youngest only currently active vent (Pu- endemic vertebrate known from Camiguin when & et nongbayan Solidum, 1985; Sajona al., 1997; we began our work (Alcala & Brown, 1998). 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, Methods though possibly somewhat earlier (Heaney 1986, 1991a, b). The island is surrounded by deep Prior Reports water, with a minimum depth to Mindanao of 385 m, which far exceeds the lowest documented Aside from a very small number of scattered late or middle Pleistocene sea lowering of ca. records (e.g., Gray 1843, who reported Paradox- 120 m (Siddall et al., 2003); thus, there is no urus hermaphroditus) and a few specimens of birds evidence of a dryland connection to Mindanao at from the 1930s deposited at the Museum of any time in the past. Comparative Zoology, the first significant surveys Camiguin Island lies within a climate zone of birds and mammals on Camiguin were characterized by annual rainfall of 2-3 m or conducted by field teams from Silliman Univer- more in the lowlands, with seasonal variation sity at 10 sites in 1967, 1968, and 1969 and on the that includes a moderately dry period from small offshore island of Mantigue in 1969. The March to May (2.0-5.5 inches/mo; ca. 5-14 cm/ field team in 1967 was led by D. P. Empesso and mo), especially wet from October to January R. B. Gonzales; we know of only bats collected by (10-15 inches/mo; ca. 25-38 cm/mo), and moist this team, all of which were deposited in the Royal during the rest of the year; mean annual rainfall Ontario Museum. Labels on specimens indicate for a 24-year period at Mambajao was that collecting was conducted at three sites, listed 99.4 inches (ca. 252.5 cm; Manalo, 1956). Tem- below as Sites 1 6—1 8. No field catalogs or notes perature declines and rainfall increases with an were kept, and no information is available on increase in elevation; based on patterns elsewhere their field procedures, but we suspect that they in the Philippines (Heaney & Regalado, 1998), followed roughly the same procedure as was used we estimate that rainfall at 1500 m is roughly in subsequent years by Dioscoro S. Rabor, as triple that at sea level (i.e., about 7.5 m/year), but follows. Field teams in 1968 and 1969, which no details from Camiguin are available. produced all the bird specimens and many of the The original vegetation of Camiguin was mammals cited here, were headed by Rabor; his lowland tropical rain forest from near the sea teams worked at seven sites, according to the to about 800 m elevation, with montane and labels on specimens in the Field Museum of mossy forest to the peaks, but by the mid-1990s, Natural History (fmnh) and Delaware Museum virtually no original vegetation remained below of Natural History (dmnh). B.R.T. Jr. worked on 300 m and little from 300 to about 800 m (Agoo, several of Rabor's teams during the 1970s and 1995). Beach and mangrove forest once occurred observed that Rabor followed a standardized along the coast, but little remains. Agoo (1995, approach to fieldwork. The field crew simulta- p. 4) described the upper elevations of Mt. neously occupied a base camp, where Rabor Hibok-hibok as a submontane plant community remained most of the time, and several satellite "dominated by shrubby plants of Radermachera, 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 were collected at two forest sites by a team led by and operate for several days to several weeks. L.R.H. and B.R.T. Jr. Our methods for sampling Members in each satellite camp would set nets mammals and birds are presented in the relevant each day. sometimes would run a small number of chapters. large snap traps, and would hunt using shotguns over an area that could readily be covered on foot. Study Sites Once each day. one member would hike to the base camp with the specimens obtained since the Camiguin is dominated by a series of volcanic and most localities refer to those last such trip, then return to the satellite camp. A peaks, peaks. similar team of collectors would operate in the We note that the three adjacent high peaks near area of the base camp as well. At the time of the center of the island (Fig. 3) are often referred to as Mt. collection, specimens were labeled simply, usually collectively Mambajao. though follow local in to the with the name of the site (often the name of we people referring northeasternmost of the three as a barangay or sitio. which could cover many major peaks Mt. and the southwesternmost as Mt. square kilometers) and rough estimate of elevation Timpoong based on topographic maps. Rabor would work Mambajao. with a team to prepare specimen labels (including Sites from 1990s measurements, date, and locality data) and to skin and stuff specimens. No field catalog or notes were We mammals at one site in 1992. usually kept: data were recorded only on the surveyed four in 1994. and two in 1995: these are referred specimen skin tags (see also Rabor. 1966). to in the their We have reconstructed the activities at Ra- species accounts by number, and locations are shown in 3. Additional bor's field sites by listing all the locality names Figure minor sites are described in the accounts. that were noted and compiling the dates on species The dates include those when which specimens were obtained. Because field only days speci- mens were obtained. Distances are teams operated simultaneously, the dates over- given from the municipal town centers, as shown in lap. We noted the elevation, when present, for Figure 2. each locality name, providing us with a range of Site 1 — Balbagon. 1 km S. 2 km E elevations for that site: many labels lack eleva- Barangay of 10 m. 12444'E tion, but we assume that the elevations noted Mambajao. 9°14.5'N, (28-29 May 1992). This site was situated in a highly apply to all specimens from a given site. We disturbed lowland area. The area believe, however, that many of the elevations as agricultural originally was lowland rain forest but in 1992 indicated on the labels of the specimens collected was a mosaic of agricultural fields, pastures, and during this period were overestimated and that small bits of secondary lowland forest. Several they should be used with caution. Additional patches of with heights of 2-4 m were comments on elevational records can be found in shrubbery present, as were several large, thick patches of specific site descriptions. erect bamboo. Six nets were set for a single night. Bats and some birds were collected. Recent Data Site 2—Barangay Balbagon. 7 km S. 2 km E r of Mambajao. 1000 m. 9 10.5'N. 124 44'E (28- Field studies that focused on mammals but 29 May 1992). A few specimens of birds were also obtained some data on birds were conducted netted in extensive, regenerating secondary forest three in during periods 1992. 1994. and 1995 by at this site. from the fmnh and Mindanao representatives Site 3—Barangay Manuyog. Sagay Munici- State Institute of University-Iligan Technology. pality. 7 km S. 3 km W of Mahinog.' 0-300 m. The first reconnaissance extended from 26 to r trip 9°5.5'N, 124 45.5'E (5-11 May 1994). Sampling 30 1992: bats were netted at a May single was conducted in heavily disturbed lowland site a field team led A. T. agricultural by by agricultural land, with very few scattered patches Peterson. The second extended from 4 period of degraded lowland forest along steep slopes 1994 to 4 June 1994: May specimens were beside rivers. Trapping and netting were con- collected at one site three forest agricultural and ducted from close to the shoreline (ca. 50 m from sites a team led by by B.R.T. Jr. The third period the beach) up to 300 m along the southwestern extended from 12 to 26 March 1995: specimens slopes of Ginsiliban Peak, extending partially

FIELDIANA: ZOOLOGY into Ginsiliban Municipality, for a total of three Site 7—Mt. Timpoong, 2 km N, 6 Vi km W of net-nights and 148 trap-nights. Mahinog, 1275 m, 9°10.5'N, 124°43.5'E (17-25 Site 4—Barangay Kital-is, Sagay Municipali- March 1995). This site was situated in primary ty, Vi km N, 6Va km W of Mahinog, 1000 m, montane forest (Fig. 3) at 1225-1350 m eleva- 9°9.5'N, 124°43.5'E (14-24 in May 1994). This tion, in the vicinity of Lasak-lasak. The average site was in disturbed transitional lowland/lower slope was ca. 35° and was often steep. The forest montane forest along the southwestern slopes of had a fairly low and relatively open canopy; the Mt. Mambajao at 900-1100 m elevation. The height of emergent trees was usually 20-25 m, slope was moderate, and the ground was covered but a few reached 30 m; dbh was 12-30 cm, and with abundant leaf litter. The vegetation con- none had buttresses. The canopy was broken by sisted mostly of small trees with dbh of 15-25 cm many treefalls, and canopy leaf sizes were small, and a few large trees standing 20-30 m high and 3-6 cm, often with serrated edges. Lower strata with 40-60 cm dbh; some lianas (5-10-cm di- leaf sizes were 4-20 cm and usually 6-10 cm. ameter), rattan {Calamus spp.), and climbing Epiphytes were abundant, including ferns, or- bamboo {Schizostachyum spp.) clung to these chids, and mosses. Arborescent pandans (Pan- emergents. Ficus spp. were common. The most clanus sp.), melastome shrubs (Melastoma spp.), common epiphytes were orchids, moss, and and tree ferns (Cyathea spp.) were common, and ferns. Tree ferns (Cyathea spp.) and rattan climbing rattan (Calamus spp.) and viney pan- (Calamus spp.) seedlings were abundant as dans (Freycinetia spp.) were abundant; Ficus and ground cover. Scattered areas of humus were Musa were rare to absent. This area had thin to up to 30 cm thick. Our total sampling effort at moderate leaf litter cover and thin to moderate this site consisted of 24 net-nights and 907 trap- humus (up to 15 cm deep); the soil consisted of nights. lightly weathered volcanic rock with many stones X Site 5— A km N, 6 !/2 km W of Mahinog, at the surface and was generally very shallow. Sagay Municipality, 1200 m, 9°9.5'N, 124°43.5'E There were no signs of human disturbance, but (26-29 May 1994). This site was situated in many trees had fallen because of very shallow disturbed lower montane forest on moderate slope soil, probably during occasional typhoons. There at 1000-1300 m elevation, in the vicinity of Lasak- were eight net-nights and 655 trap-nights. lasak (Site 12). There was some evidence of small- Site 8—Mt. Timpoong, 2% km N, 6 '/a km W scale illegal logging done about a year previously. of Mahinog, 1475 m, 9°11'N, 124°43.5'E (22-25 Emergent trees had dbh of between 40 and 60 cm March 1995). Sampling at this site was con- and heights up to 30 m. Lianas and canopy ducted in mossy forest. The site was on a steep epiphytes (mainly ferns and mosses) were present. slope, averaging 50°, and ranged from 30-70°, Rattan {Calamus spp.) and climbing bamboo including steep gullies. The height of the canopy {Schizostachyum spp.) were also present, with was usually 8-10 m but varied from 2 to 3 m in pitcher plants {Nepenthes) present but rare. Un- exposed spots to a maximum of ca. 18 m in low, derstory and ground cover consisted of rattan protected places; trees were generally gnarled, seedlings and ferns, with some sedges. Leaf litter with dbh of 12-20 cm (rarely 25 cm). Canopy covered about 80% of the ground, and humus was leaves were small, with serrated edges, typically typically about 30 cm thick. Our total sampling 1-8 cm, but most were 4-5 cm. Oaks, laurels, effort consisted of 339 trap-nights; at this site, nets tree ferns, and arborescent Pandanus were were concurrently maintained with those at Site 6 common. Lower-strata leaves were small, usually for a total of 14 net-nights. 4-8 cm. Ficus and Musa were rare to absent. Site 6—Barangay Kital-is, Sagay Municipali- Epiphytes were abundant, including mosses, ty, on a small peak near Mt. Mambajao, 1 km N, ferns, orchids, and saplings, and pitcher plants 7'/2km W of Mahinog, 1300m, 9°10'N, (Nepenthes spp.) were common. Canopy vines 124°43'E (26-29 May 1994). This site was (Freycinetia spp. and Calamus spp.) were abun- situated in primary mossy forest at 1200- dant. Ground plants included ferns, saplings, 1400 m. Hanging moss was abundant on trees, and abundant pandans (Freycinetia). The ground but moss cover on the ground was light and the was covered by abundant and thick leaf litter humus layer not more than a few centimeters over deep humus (over a half meter thick, with thick. Our total sampling effort consisted of 348 many tunnels and vacuities). Moss covered trap-nights for this site and 14 net-nights for nearly all tree trunks, branches, and fallen logs Sites 5 and 6 combined. and hung from branches in sheets and often

HEANEY AND TABARANZA: MAMMAL AND LAND BIRD STUDIES cited the team to have been covered the ground. No human disturbance was specimens by seen, but there was evidence of some large collected on this mountain above this elevation landslides and many tree falls. No netting was were probably obtained close to the peak. — Catarman performed at this site; there were 386 trap- Site 10 Gidag-on, Municipality, ft 1 50-450 nights. 500-1500 (ca. m) [approx. 9T0.5'N, 124°39.5'E] (13-28 June 1968). This was the Sites from the 1960s lowest campsite during the 1968 field season; birds and mammals were collected. Birds were

As noted above (see Methods), field teams collected mostly from 23 to 28 June 1968, with from Silliman University conducted surveys on a few taken on earlier dates (13, 16-17, and 19 collected on 13 Camiguin at 10 sites in 1967-1969 plus an June). Some birds June were additional site on the small, adjacent Mantigue noted to have been collected at 2000 ft (ca. Island. As noted above (Methods), we have 600 m), possibly the highest point reached from estimated the location of these sites; our esti- this site. Because it was the lowest campsite in mates are shown in brackets. The vegetation 1968, it is the most likely location of the base types and condition at these sites were not noted camp that year. We are unable to locate this site, by the collectors. We surmise, on the basis of but some collecting dates at this site overlapped specimens collected here and the condition of the with Sites 9 and 1 1; we suspect that it was on the remaining vegetation on Camiguin in the 1990s, lower slopes of Mt. Catarman. No information that when the collecting was done, the vegetation on the vegetation when the specimens were below 800 m ranged from secondary lowland collected at this site is available, but we surmise forest to heavily populated agricultural areas. By that it would have been partly secondary lowland the time of our surveys in the 1990s, the lowland forest and partly agricultural. forest was almost totally cleared and replaced Site 11 —Kasangsangan, Catarman Munici- with coconuts, and hardly any area below 800 m pality, 1000-2500 ft (300-800 m) [approx. supported remnant forest except along steep 9 ll'N, 12440' E] (11-22 June 1968). This was slopes. During the 1960s, from ca. 1000 m up to the middle campsite in 1968. We are unable to the peak, the sites would have been covered with locate this site, but because collecting dates primary montane and mossy forest, as they were overlapped with Site 9, we suspect that it was in the 1990s. mainly on the lower to middle slopes of Mt. Site 9—Mt. Catarman, Catarman Municipal- Catarman. Several specimens were collected up

ity, 2000-4950 ft (ca. 600-1500 m) [approx. to 4950 ft (ca. 1500 m), which might indicate 5.5 km S, 4.5 km W Mambajao, 9 12'N, that the team assigned to this site ventured all the 124 40.5' E] (10 29 June 1968). This was the way to the peak of Mt. Catarman, but we suspect highest site surveyed in 1968; both birds and mixing of localities, as noted above. No in- mammals were collected. Mammals were collect- formation on the vegetation when the specimens ed at 2500-4500 ft (ca. 750-1400 m). The ma- were collected is available, but we surmise that, jority of the birds specimens were taken on 12-21 similar to Site 9, it would have been partly June, but a few others were collected on 10, 25, secondary lowland forest and partly agricultural. and 29 June. Bird specimens were taken from Site 12—Mt. Timpoong, Lasak-lasak, Mahi- 2000 to 4950 ft (ca. 600-1500 m). This would nog Municipality, 4400-5700 ft (ca. 1350- suggest that the team sent to this camp concen- 1700m) [approx. 9 1 l'N, 12443. 5'E] (19-28 trated their efforts from the middle to the upper June 1969). We traveled through this site in slopes, including the peak of Mt. Catarman, but 1995; it is located near the headwaters of the several were taken at 1000 ft (ca. 300 m) on 14 Sagay River, from 1200 m to roughly the peak. and 16 June 1968. However, we also know that This was the highest site surveyed in 1969; birds collecting at this site was done simultaneously and mammals were collected. Collecting activi-

with Site 1 1 , and specimens from both sites were ties were conducted from 19 to 28 June. The then taken down to the base camp at Site 10 for majority of the bird specimens were taken at processing by D. S. Rabor. We suspect that in 4800-5700 ft (ca. 1500-1700 m), indicating that the there process, was some mixing up of the team concentrated their collecting in the elevational data for some specimens (see also high-elevation habitats on Mt. Timpoong. Sev-

Site 1 1 ). the the Furthermore, highest peak of eral specimens were taken slightly farther down, Mt. Catarman does not exceed 1400 m, so all the at 4400 ft (ca. 1350 m). On 23 June 1969, some

FIELDIANA: ZOOLOGY specimens attributed to this site were noted as this site. Mt. Mambajao's highest peak is ca. taken at 800 m. As with the previous year at Sites 1600 m; this location in Catarman Municipality 9-11, collecting at this site was done simulta- suggests that specimens were taken from the neously with Site 13, and specimens were taken middle slopes, on the southern to southwestern down to the base camp at Site 14 for processing. flank of the mountain. We suspect that some mixing up of elevational Site 17—Mt. Mambajao, Sangsangan, Catar- data for some specimens from among the three man Municipality, 1400-3300 ft (ca. 400- sites occurred. While the vegetation at this site was 1000 m) [approx. 9°9'N, 124°42.5'E] (14-31 not indicated when the 1960s collecting was May, 15 June 1967). Only mammals were undertaken, it would have been primary mossy collected at this site. This location suggests that forest, based on the condition of the habitat that specimens were taken on the lower to middle we saw on Mt. Timpoong in the 1990s. slopes of Mt. Mambajao, on the southern to Site 13—Mt. Timpoong, Matugnao, Mahinog southwestern flank of the mountain. Municipality, 3150 ft(ca. 950 m) [approx. 9°10'N, Site 18—Tag-ibo Cave, Catarman Municipal- 124°44.5'E] (12-26 June 1969). This was the ity, 400 ft (ca. 100 m) (31 May 1967). A few bats middle campsite established in 1969; birds and were collected at this site, which we have been mammals were collected. The majority of the bird unable to locate. Because it was visited on the specimens bear the elevation 3150 ft (ca. 950 m), same day as specimens were obtained from Site with a few at 3250 ft (ca. 1000 m), around the 17, they are probably near each other. midslopes of Mt. Timpoong. A few specimens, Site 19—Mantigue Island [approx. 2 km N, however, were noted as taken at 800 ft (ca. 250 m) 4 km E Mahinog; 9°10.5'N, 124°49.5'E] (28 June on 1 6 June and at 4800 ft (ca. 1 500 m) on 20 June. 1969). This site is a 4-ha coralline island, situated Collecting at this site was done simultaneously ca. 3 km east of Barangay Hubangon, Mahinog with Sites 12 and 14, and we suspect that some Municipality (Figs. 2 and 3). It was visited on 28 mixing of elevational data occurred. June 1969, and only White-collared Kingfisher Site 14—Puntod, Mahinog Municipality, {Halcyon cloris), Yellow-vented Bulbul {Pycno- 800 ft (ca. 250 m) [approx. 2 km N, 2 km W notus goiavier), Pied Triller (Lalage nigra), and Mahinog; 9°9.5'N, 124°46'E] (24-29 June 1969). Olive-backed Sunbird {Nectaruna jugularis) were This was the lowest campsite established in 1969; collected. The vegetation when it was visited in birds and mammals were collected. Bird collect- 1969 was most likely beach forest, with some ing took place on 24-29 June. Elevations were all disturbance. given as 800 ft (ca. 250 m), with a single bird specimen bearing an elevation of 700 ft (ca. Conservation 200 m). Puntod is a village located on the lower slopes of Mt. Timpoong, about 2 km west of the As documented by Balete et al. (2006), Heaney town center of Mahinog (Fig. 3). This probably et al. (2006), and Tello et al. (2006), Camiguin was the base camp during the 1969 field season. supports at least 24 species of mammals and at The vegetation in the 1960s is unknown, but least 54 species of birds, and additional fieldwork given its situation as a settled village during that is likely to document the presence of additional time, the habitat was probably already largely species. This includes two species of mammals (in agricultural with remnant lowland forest. the genera Apomys and Bullimus) and one species Site 15—Mount Timpoong Peak, Mahinog of bird (genus Loriculus) that are unique to Municipality, 5700 ft (ca. 1600 m) [approx. Camiguin; Camiguin is the smallest island in the 9°11'N, 124°43.5'E]. Only a single specimen of Philippines currently known to support unique Simcus murinus bears this locality. Given the species of mammals (Heaney, 1986; Heaney et restriction to Mahinog Municipality, the site al., 1998) and is smaller than any island pre- must have been at the peak on the eastern side of viously known to support an endemic bird the mountain, near to or including Site 8. The (Peterson et al., 2000). The endemic species of vegetation at this site would have been mossy mammals have been documented to occur in forest, as described for Site 8. lowland, montane, and mossy forest from Site 16—Mt. Mambajao, Mahidlaw, Catar- 1000 m to near the peaks but probably occur at man Municipality, 2500-3500 ft (ca. 800- lower elevations as well where we were unable to 1000 m) [approx. 9 9.5'N, 124°42.5'E] (24 and sample; the endemic bird (Tello et al., 2005) has 26 May 1967). Only a few bats were collected at been documented from ca. 300 to 1200 m, which

HEANEY AND TABARANZA: MAMMAL AND LAND BIRD STUDIES encompasses lowland and lower montane forest. trees were often killed deliberately by the loggers The presence of these unique species, along with so that they could then be cut down. Slash-and- the frog and plants that are restricted to the burn agriculture {kaingin) of the remnant forest island, have caused Camiguin to be listed as a key usually followed quickly (Heaney & Tabaranza. national and global priority site for conservation 1997). By the time we visited the island in 1995. (Mallari et al., 2001: Heaney & Mallari. 2002: areas below 300 m had no native forest, and only Ong et al.. 2002) that was overlooked by a little second growth remained up to ca. 800 m. previous assessments that were based on in- Only the montane and mossy forest was in complete biological surveys (e.g.. Hauge et al.. relatively good condition, but small-scale logging 1986: Heaney. 1993: Peterson et al., 2000). and kaingin was creeping farther up on the These diverse mammal and bird faunas mountains: we noted a new kaingin at 1200 m on originally occurred along the entire elevational the side of Mt. Timpoong in 1995 (Heaney & and habitat gradient. The most fundamental Tabaranza. 1997: Mallari et al.. 2000). requirement for their conservation is the contin- It is essential to note that conservation of the ued existence of substantial areas of all original forest will have great benefits to the people of types of habitat in mature, good condition. The Camiguin as well as to the wildlife. The forest areas must be large enough for the existence of can serve as a permanent source of wood and substantial populations, not just a few individu- other nontimber forest products to the local als, since populations often are vulnerable to inhabitants if this is carefully managed. Perhaps extinction when they drop below about 1000 most crucially, the forests of Camiguin provide individuals (Primack. 1998). the people with abundant water for domestic, Fortunately. Camiguin retains enough forest agricultural, and industrial uses. As an addition- cover to make this possible, though there are al essential ecological service, the forest helps a number of challenges to surmount. The results prevent soil erosion and landslides. In early of the Swedish Satellite Survey of forest cover in November 2002. about 200 human lives were lost 1987 (National Mapping and Resource Informa- and thousands of houses and other properties tion Authority. 1988) showed a fairly large area damaged by landslides during a typhoon, partic- of forest on Camiguin. mostly at upper eleva- ularly in Mahinog. This disaster demonstrates tions on Mt. Mambajao and Mt. Timpoong and the great economic and human cost of forest associated highlands, as shown in Figure 2. Our degradation. Additionally, the continued popular- surveys in 1994 and 1995 showed that much of ity of Camiguin as a tourist destination will rely that forest still remained, though the edges had strongly on its ability to provide beautiful scenery, been further degraded. Downslope. the degree of attractive areas for hiking, fresh and clean water, disturbance progressively increased so that little and health} coral reefs (which require low levels of if any lowland forest remained in a primary siltation from adjacent rivers). condition: indeed, most had been replaced with The importance of protecting the remaining coconut plantations, agricultural areas, and forest of Camiguin can be appreciated when one grassland. In 2001. it was estimated that S29c considers its fast-growing population. In 1980, the of the island, ca. 20.847 ha. was covered by population of Camiguin was about 57.000; two croplands and other inhabited areas: almost decades later, in 2000. the population had in- half the island's area was covered with coco- creased to more than 73.000: in 2020. it is projected nut plantations (http://agrilO.norminet.org.ph/ to increase to 90.000 (http://www.popcom.gov. NMProfile/profile_camiguin.htm). leaving \S9c ph/sppr/statistics/Ieastvis_wesmin_northmin.htm). as secondary and primary forest. Consequently, the population density of Cami- 2 According to local residents, much of the guin increased from 279 persons/km in 1980 to forest on Camiguin was removed by commercial 306 persons/km" in 2000: these were well above 2 logging operations in the 1980s, with the logged the national density of 160 persons/km and 2 areas subsequently being cleared for agriculture. 251 persons/km in 1980 and 2000. respective- During the 1990s, small-scale commercial log- ly (http://www.popcom.gov.ph/sppr/statistics/ ging, operating under salvage permits, was the lleastvisz_wesmin_northmin.htm: http://www. main agent responsible for the continuing de- popcorn. gov.ph/sppr/statistics/table3. htm). nudation of the areas below 1000 m. These In response to all these issues, including our salvage permits technically allowed only dead discoveries in 1994-1995 of endemic species of trees to be cut. but local residents told us that mammals, the local gov ernment and Department

10 FIELDIANA: ZOOLOGY of Environment and Natural Resources (DENR) elevation. This will enable future management to have moved steadily in recent years to have the rehabilitate/restore lowland forest, which is one upland areas where forest remains declared the of the most critical habitats on Camiguin. "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 Acknowledgments and is expected to soon be endorsed to the For assistance with the often field president. If it is successful at that level, as is challenging work on we thank N. E. anticipated, it must then be voted on by the Camiguin, Antoque, N. N. M. R. Philippine Congress for final designation, a pro- Batara, Batocael, Bojo, Carmona, A. M. L. cess that may yet require several years. The Fernandez, DeOcampo, Jayoma, approximate proposed boundary of the natural Mostrales, A. T. Peterson, G. Rosell, A. Tabar- B. Tabaranza and D. Tabaranza. monument, with an area of 2,228 ha, plus anza, III, Permits were the Protected Areas a 1,423-ha buffer zone, is shown in Figure 2. provided by Although both the location and area of forest and Wildlife Bureau of the Department of Environment and Natural Resources and the natural monument as shown in Figure 2 (DENR); we offer thanks to A. C. C. are approximate, it is clear that the majority of special Alcala, M. and Pollisco and to the primary forest lies within the proposed Custodio, Mendoza, W. the 10 Office of the DENR. Emmauel boundaries of the park. Region Aranas and Primitivo essential We fully endorse these recommendations and Espinas provided assistance and on actions. But further, as part of this program of logistical support Camiguin the fieldwork. We thank E. C. environmental protection and stabilization, we during Canete, and G. Rosell-Ambal for information recommend the following to the local govern- Custodio, on the status of the area. For the loan of ment of Camiguin and the Philippine DENR as protected under their care, we are indebted to M. essential activities: 1) continue and expand active specimens Carleton and H. Kafka and P. enforcement efforts to protect existing forests, (usnm) Myers we thank G. Hess for wildlife, and environment; 2) continue and (ummz); especially (dmnh) much assistance and over expand current reforestation projects, using only encouragement many The and were native species of trees, not exotics, because the years. maps graphs prepared by Lisa the of was exotic trees do not provide habitat for the Kanellos; photograph Camiguin Rebecca Banasiak. We biodiversity of Camiguin and are injurious to digitally prepared by thank P. for information on the the soil; 3) cancel existing salvage cutting permits Janney geological for dead and fallen trees because these have history of Camiguin and J. Bates, N. Collar, and E. Rickart for constructive comments on an often been abused; and 4) complete the process earlier draft of the This research of declaring the remaining forest and key manuscript. was the World Environment and watershed areas a national protected area as supported by Resources of the John D. and Catherine soon as possible. To the maximum extent Program T. MacArthur Foundation and the Marshall possible, all portions of the protected area by Field Fund, the Ellen Thorne Smith Fund, and should be connected by corridors of mature (or the Barbara Brown Fund for Mammal Research regenerating) habitat, especially along rivers and of the Field of Natural streams. Museum History. In connection with the above recommenda-

tions, it is essential to both wildlife conservation and watershed that national and local protection Literature Cited governments include substantial lowland areas (those below 800 m) in the proposed national Agoo, M. G. 1995. Camiguin Island. Philippine park, regardless of the present condition of the Biodiversity Information Center Plant Unit News- letter, 3: 4. habitats and vegetation, especially including all A. and W. C. Brown. 1998. areas within 200 m of streams and rivers. We Alcala, C, Philippine Amphibians: An Illustrated Guide. Bookmark, therefore recommend that the natural monument Makati City. be expanded to include all good-quality second- Balete, D. S., B. R. Tabaranza, Jr., and L. R. ary forest at all elevations and to include all Heaney. 2006. An annotated checklist of the land major watercourses down to at least 600 m of birds of Camiguin Island, Philippines, pp. 58-72. In

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12 FIELDIANA: ZOOLOGY Primack, R. B. 1998. Essentials of Conservation SlDDALL, M., E. J. ROHLING, A. AlMOGI-LaBIN, C. Biology. Sinauer Associates, Sunderland. HEMLEBEN, D. MEISCHNER, I. SCHMELZER, AND D. A. Smeed. 2003. Sea-level fluctuations the last PUNONGBAYAN, R. S., AND R. U. SoLIDUM. 1985. during 423: 853-858. Camiguin Island. Philippine Institute of Volcanolo- glacial cycle. Nature, gy and Seismology, Quezon City. Stattersfield, A. J. M., J. Crosby, A. J. Long, and Rabor, D. S. 1966. A report on the zoological D. C. Wege. 1998. Endemic Bird Areas of the Priorities for expeditions in the Philippines for the period 1961 World: Conservation. Birdlife Interna- 1966. Silliman Journal, 13: 604-616. tional, Cambridge. Rickart, E. A., L. R. Heaney, and B. R. Tabaranza, Tello, J. G., J. F. Degner, J. M. Bates, and Jr. 2002. Review of Bullimus (Muridae: Murinae) and D. E. Willard. 2006. A new species of Hang- description of a new species from Camiguin Island, ing-Parrot (Aves: Psittacidae: Loriculus) from Philippines. Journal of Mammalogy, 83: 421-436. Camiguin Island, Philippines, pp. 49-57. In Heaney, L. The and Birds of Cami- Sajona, F. G., H. Bellon, R. C. Maury, M. R., ed., Mammals a Distinctive Center of PUBELLIER, R. D. QUEBRAL, J. COTTEN, F. E. BAYON, guin Island, Philippines, Fieldiana 106: 1- E. Pagado, and P. Pamatian. 1997. Tertiary and Biodiversity. Zoology, n.s., 72. Quaternary magmatism in Mindanao and Leyte (Philippines): Geochronology, geochemistry, and Wildlife Conservation Society of the Philippines tectonic setting. Journal of Asian Earth Sciences, 1997. Philippine Red Data Book. Bookmark, 15: 121-153. 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 2 Lawrence R. Heaney 1 and Bias R. Tabaranza, Jr.

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 concentrations of unique mammalian diversity worldwide (Heaney et al., 1998). These species The Philippine Islands are notable for their are usually not widely distributed within the large number of unique species of mammals; of Philippines but rather are confined to one or 172 species known in 1998, 111 (64%) occurred a few islands. Recent studies have shown that the nowhere else in the world, one of the greatest geological history of the archipelago is largely responsible for the pattern of distribution, with 1 Field most of found on one of Museum of Natural History, 1400 South species mammals only Lake Shore Drive, Chicago, IL 60605-2496, U.S.A. the several islands that formed during periods of 2 low sea level in the late and middle Pleistocene. Department of Biology, Iligan Institute of Tech- Mindanao State Each nology, University, Iligan City, of these Pleistocene islands is surrounded Lanao del Norte, Philippines. 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 fmnh 147874-876, 147880, 147900-904, 147906, oceanic island throughout its existence. Howev- 148125-128, 148132, 148135-138. Apomys in- er, though they are isolated by deep-water signis—Mindanao Island: Bukidnon Province: channels, the channels are not wide, usually not Mt. Kitanglad Range: 16.5 km S, 4 km E Camp more than 25 km and often much narrower Phillips, elev. 1900 m, fmnh 148152; 17 km S, (Heaney, 1986, 1991, 1993, 2004; Heaney and 7 km E Baungon, elev. 1550 m, fmnh 146703; Rickart, 1990; Heaney & Regalado, 1998). 18 km S, 7 km E Baungon, elev. 1800 m, fmnh Camiguin Island was noted by Heaney (1984, 146704-710, 146712-714, 146716-718, 147088- 1986) as an apparent exception to this pattern: 089, 147091-092,— 147094, 147098-099, 147102. collections made on Camiguin in the 1960s by Apomys sp. Biliran Island: 3 Vi km S, 5 Vi km field teams from Silliman University did not W Caibiran, elev. 700 m, ummz 160314, 160290, include any endemic mammals, even though the 160429^130; 4'/2 km S, 5 km W Caibiran, elev., island seemed large enough to support them 920 m, ummz 160290-291, 160315-316. Leyte (Heaney, 1986, 2004). After the discovery of four Island: 9 km N, 3 km E Baybay, elev. 750 m, endemic species on Sibuyan Island, another small ummz 160318; 8 !/2 km N, 2'/2 km E Baybay, oceanic island in the & elev. 500 ummz 160441. n. archipelago (Goodman — m, 160317, Apomys Ingle, 1993; Heaney et al., 1998), we suspected sp. Holotype and referred specimens from that the mammals of Camiguin might not have Camiguin Island (see below). been fully surveyed, so we returned in 1994 and Specimens were assigned to age categories as 1995 for further investigations (Heaney et al., follows. Subadult animals are those that have 2004; Heaney & Tabaranza, 2006). In the course not completed cranial growth, especially those of those field studies, we documented the presence having unfused basicranial sutures; these young of two previously unknown species of mammals animals have pelage that is usually softer and (Heaney & Tabaranza, 1997), Bullimus gamay grayer than that of adults and are noticeably (Rickart et al., 2002) and a species of forest lower in weight, and females are usually nullip- mouse, genus Apomys. It is the purpose of this arous. Young adults are older; they have unworn paper to describe this new species of forest mouse. 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- Materials and Methods ed cranial growth and have adult pelage, and usually the females are multiparous. Terminolo- Specimens examined for this study are housed gy for description of external features follows in the Delaware Museum of Natural History Brown (1971) and Brown and Yalden (1973). (dmnh), Field Museum of Natural History Terminology for cranial and dental features (fmnh), Mindanao State University-Iligan In- follows Musser and Heaney (1992). Scanning stitute of Technology (msu-iit), National Muse- electron micrographs of skulls and teeth were um of the Philippines (nmp), University of made with uncoated specimens. Michigan Museum of Zoology (ummz), and External measurements (total length, tail States National of Natural United Museum length, hind foot length, length of ear from History (usnm). Half the specimens from Cami- notch, weight in grams) were taken from in will nmp. guin now housed fmnh be sent to collectors' field catalogs or specimen labels. Material examined included specimens prepared Fourteen cranial measurements (Table 1) were skins skulls with as study with (and some taken with digital calipers to the nearest 0.01 mm postcranial skeletons), complete skeletons, and by Heaney; comparisons made in the text refer formalin-fixed specimens stored in 70% ethyl only to specimens also measured by Heaney. alcohol, many with skulls subsequently removed and cleaned. The following samples were exam- ined: Apomys hylocoetes—Mindanao Island: Bukidnon Province: Mt. Kitanglad Range: Results 15 km S, 12.5 km W Dalwangan, elev. 2800 m, fmnh 148055; 16.5 km S, 4 km E Camp Phillips, The endemic Philippine genus Apomys was elev. 1900 m, fmnh 147871-872, 148123-124; described by Mearns (1905) to contain three A. 18.5 km S, 4 km E camp Phillips, elev. 2250 m, species: A. hylocoetes (as the type species),

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE 15 = "3 insignis, and A. petraeus (the last synonymized A. "A/C" (Negros) with A. hylocoetes by Musser, 1982), all from Mt.

from the I Apo, which name was derived. As rC"A "B" (Sibuyan) related by Musser (1982), additional species were named from Luzon and Catanduanes, and the A. musculus genus was found throughout much of the archipelago (Ruedas, 1995; Heaney et al., A. microdon 1998). However, because the initial description was vague and many genera of Indo-Australian A. hylocoetes rodents poorly studied, Apomys was briefly with the Rattus at synonymized genus a time A. insignis when most Indo-Australian murids were placed in that genus. Musser (1982) thoroughly rede- A. "F" (Leyte) scribed Apomys, pointing out its many distinctive and redefined the then characters, species A. camiguinensis known. Musser and Heaney (1992) further defined and compared Apomys to other Philip- A. datae pine murids, and they pointed out its apparent close relationship to Chrotomys, Celaenomys, r A. gracilirostris and Rhynchomys, also endemic to the Philip- pines. Using data from Musser and Heaney Rhynchomys isarogensis (1992), Heaney and Rickart (1990) postulated that Apomys was basal to the clade including c Chrotomys gonzalesi Chrotomys, Celaenomys, and Rhynchomys and noted the diversification of this clade within the Fig. 1. Hypothesis of phylogenetic relationships within the genus Apomys based on parsimony analysis Philippines as an example of adaptive radiation. of cytochrome-b molecular data (based on Steppan et et al. (1998) noted the of Heaney presence many al., 2003). undescribed species of Apomys, including the species from Camiguin reported by Heaney and Tabaranza (1997). infraorbital branch of the is Rickart and Heaney (2002) showed that Ap- stapedial artery] and of the is omys hylocoetes, A. insignis, and the Leyte Apomys partially open part artery exposed on the ventral surface of each (as well as most others from Greater Luzon and pterygoid plate. That combined with Greater Negros-Panay) have distinctive karyo- feature, bright tawny brown or types, but the Apomys from Camiguin has not upperparts, buffy underparts, pale and on the tail been karyotyped. Steppan et al. (2003) used buffy feet, patterning (usually monocolored or bi- molecular data to assess phylogenetic relation- mottled, rarely sharply the five in ships and geographic patterns of diversification colored) suggest species [known be more related to each other within Apomys. Analysis of variation in cyto- 1982] may closely than to others in Musser's chrome^ in 10 species (Fig. 1) showed the any Apomys." (1982) of this based presence of three major clades: one containing recognition and diagnosis clade, on few was and Apomys datae and A. gracilirostris; a second clade very specimens, prescient In containing A. microdon, A. musculus, and two remains accurate. the analysis by Steppan al. the is basal undescribed species from Negros and Sibuyan (the et (2003), species from Camiguin "Greater Luzon, Mindoro, and Negros clade"); to the others in the "Greater Mindanao clade" and a third clade containing A. hylocoetes and A. (Fig. 1), and they estimated the time of of the the insignis from Mindanao, plus an undescribed divergence Camiguin mouse from its million species from Leyte, Biliran, and Bohol and other members of clade at about 2.3 ± about 25%. another from Camiguin ( the "Greater Mindanao years clade"). The second and third of these three clades We note that the Apomys from Biliran, Bohol, form a monophyletic clade that Musser (1982) and Leyte (which we hereafter refer to as "the described as being "species ... of small or Leyte Apomys''') was tentatively considered to medium body size in which the canal for the represent either A. microdon (by Musser, 1982) internal maxillary artery [also described as the 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 Mt. Timpoong, Camiguin Island, in March 1995.

1998). but the combination of the morphological and cleaned. Specimen is currently on deposit at data presented here, karyotypic data (Rickart & fmnh and is to be transferred to nmp. Heaney, 2002). and molecular data (Steppan et Type Locality—Barangay Kital-is, Sagay Mu- al., 2003) have led us to conclude that the Leyte nicipality, 2 km N, 62 km W Mahinog, 1000 m Apomys is a distinct species; further details and elevation, Camiguin Province, Camiguin Island. description will be published elsewhere. 9°9.5'N, 124°43.5'E (see Heaney & Tabaranza, The specimens from Camiguin are morpho- 2006, for further details). logically similar to series of A. hylocoetes and A. Referred Specimens and Localities—In addi- insignis from Mindanao and the undescribed tion to the holotype, 19 paratypes are known species from Leyte and Biliran but readily from three localities ranging from 1000 to distinguished from all three of these species on 1400 m (fmnh 154815-154816, 154854-154860, the basis of external and cranial features. Heaney 167878-167882, plus 6 specimens at msu-iit); for et al. (1998) and Steppan et al. (2003) referred to localities, see Heaney et al. (2006). All were this animal informally as "'Apomys sp. D." We originally preserved in formalin and are now now name the species from Camiguin as Apomys stored in ethyl alcohol, many with skulls re- camiguinensis, new species. moved and cleaned, or were prepared as complete skeletons. Tissue samples are housed at fmnh. Half of the specimens will be deposited at nmp. new Apomys camiguinensis, species Distribution—Known only from the upper elevations on Mt. Timpoong, Camiguin Island, Adult male, fmnh Holotype 167878, collect- but probably occurring throughout the montane ed 16 1994 B. R. May by Tabaranza, Jr. and mossy rain forest on Camiguin Island (see Specimen fixed in trans- originally formalin, fig. 2 in Heaney et al., 2004) and possibly at ferred to im ethyl alcohol with skull removed lower elevations.

IS FIELDIANA: ZOOLOGY Measurements—Table 1. rolabial cusp of the third upper molar is barely Etymology—The specific name refers to the evident in most individuals. sole island on which the species is found. We Description and Comparisons—Apomys cami- suggest the common English name "Camiguin guinensis is an attractive mouse with large eyes forest mouse." — and ears, long tail, and soft pelage (see Frontis- Diagnosis A species of the genus Apomys, as piece, this volume, and Fig. 2). As with other defined by Musser (1982) and Musser and members of the genus, the pelage is soft and Heaney (1992), including the following distinc- dense, without spines or stiff hairs. The dorsal tive generic features: rostrum long and moder- coloration is a rich brownish-russet with a small ately narrow; viewed laterally, rostrum with amount of salt-and-pepper speckling; underfur is a rectangular shape, with premaxillaries project- pale slate-gray. The venter is paler, usually ing well anterior to the anterior edge of the nearly white with a wash of buffy or pale russet, upper incisors; incisive foramina broad relative but some individuals have blazes of pure white to length; bony palate wide and long, densely (usually on the chest) or are much darker brown pitted and perforated; upper third molar re- or russet-brown. There is a narrow area of bare duced to a large round peg; lower third molar skin around the eyes; the ears are moderately also peg-like but retaining an anterior lamina, dark brown, with short hairs apparent on the without evidence of the two cusps that usually outer surface, and present but tiny and nearly form this lamina; occlusal surface of each first invisible on the inner surface. The mystacial and second upper molar consisting of two simple vibrissae are long and conspicuous. The dorsal chevron-shaped laminae followed by a small oval surface of the fore and hind feet are mostly buffy lamina, without evidence of cuspidation; audito- or pale brown but with a narrow band of ry bulla separated from the squamosal and scattered darker hairs around the midline, and alisphenoid by a gap that is formed by the these decrease in number and length toward the coalescence of the postglenoid foramen, the distal end of each foot (with only pale hairs on postalar fissure, and the middle lacerate fora- the dorsal surface of the toes). The feet are lightly men. pigmented or unpigmented on the ventral As described in more detail in the following surface, with conspicuous plantar pads on the section, the Camiguin mouse is defined by the ventral surface (Fig. 3). The tail is long with following characters or unique combination of conspicuous scales; fine hairs that are present characters: moderate body size but somewhat between the scales are most visible on the dorsal robust build for the genus overall; the tail is long and lateral surfaces and least visible ventrally. relative to body length, with unusually large The tail is darker on the dorsal surface than on scales, and more often with a sharp transition the ventral surface. The scrotum of adult males is from dark brown dorsum to pale brown venter fairly small and projects beyond the abdomen

(i.e., sharply bicolored) than in other species; only partially on the posterior portion and has moderately long but unusually broad hind foot black or dark brown pigment at the posterior tip, with small plantar pads, and slightly shorter ear. about 3-5 mm in length. Females have two pairs The pelage has more brown in the generally of inguinal mammae. russet-brown dorsum than on its closest relatives Apomys camiguinensis is easily distinguished and has more conspicuous salt-and-pepper from most members of the genus by its in- speckling dorsally; the mystacial and genal termediate size (only A. insignis and A. hylo- vibrissae are long but moderate for the genus. coetes are similar) and from those two species by The cranium has an unusually long braincase both external and cranial characters. Apomys and orbit, somewhat narrow zygomatic and camiguinensis has total length (average 254— mastoid width, and a moderately long but deep 260 mm) slightly greater than A. hylocoetes and robust rostrum. The palate is rather narrow, (248-252 mm) and A. insignis (251-252 mm) the posterior edge of the palate extends un- and substantially greater than the Leyte Apomys usually far posterior to the last molar, the (238-245 mm; Table 1). The average tail length incisive foramina long and wide, and the distance (146-148) is equal to that of A. insignis (147— from the posterior edge of the incisive foramina 148 mm), and substantially longer than that of is unusually short. The longest axis of the bullae A. hylocoetes (141-142 mm) and the Leyte is about 35° from the midline axis of the skull. Apomys (140-145 mm). The tail averages 57% The toothrows are of moderate size; the ante- of the total length in A. camiguinensis, compared

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE 19 A. Fig. 3. Ventral surface of the right hind feet of (A) Apomys camiguinensis, (B) A. hylocoetes, (C) 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 more of the salt-and-pepper appearance; the 59% in the Leyte Apomys. The hind foot (about Leyte mouse is dorsally brighter red than the 33 mm) is about equal in length to that of A. others, with more red and orange in the russet insignis (33 mm) and is substantially longer than than the other three and almost no salt-and- in A. hylocoetes (31-32 mm) and the Leyte pepper. Ventral coloration is generally similar in Apomys (30-31 mm); the hind foot of A. all four, though with the variation noted above, hylocoetes is notably the broadest (Fig. 3; see but A. hylocoetes tends to have more of an also fig. 7 in Musser, 1982). Ear length is greatest orange wash than the others. The dorsal surface in A. hylocoetes (20.2 mm), with A. insignis of the hind feet usually is palest in the Leyte (19.5 mm), A. camiguinensis (19.0 mm), and the mouse and darkest in the Camiguin mouse. The Leyte Apomys (18-19 mm) progressively slightly mystacial vibrissae are long on all four species smaller 1 (Table ). The weight of A. camiguinensis but longest on A. insignis (up to 56-60 mm (38-41 g) averages the greatest of the four, maximum), on which they reach past the middle followed by A. hylocoetes (36-39 g), A. insignis of the back, and intermediate on the Camiguin 38 (37 g), and the Leyte Apomys (28-31 g). In mouse (52-55 mm), A. hylocoetes (51-55 mm), other words, the Camiguin Apomys is relatively and Leyte mouse (50-55 mm). Genal vibrissae and heavy long and has a relatively long tail (but reach to the anterior edge of the largest lateral slightly less long proportionately than some close pad on the Camiguin and Leyte mice but farther a relatives), moderately long hind foot, and forward, to the base of the toes, on A. hylocoetes somewhat short ear. and A. insignis. The external following qualitative characters The hind foot (Fig. 3) differs markedly among also distinguish Apomys camiguinensis from its the four. The foot of A. camiguinensis is about three closest relatives. The dorsal coloration of the same length as that of A. hylocoetes but is the mouse is less Camiguin russet and more broader and has smaller plantar pads. The hind brown than in A. and A. hylocoetes insignis, with 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 eoetes are slightly thinner and more sharply pads of moderate size. On all species, the claws pointed and the digits bearing them slightly more on all five digits are unpigmented and laterally slender. The digits and claws of A. insignis are compressed. On A. eamiguinensis, the claws 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 camiguinensis being the most robust, A. hylo- slender. The forefeet of A. camiguinensis have coetes slightly shorter and thinner, A. insignis smaller pads than those of the others (but only much shorter and more slender, and the Leyte proportionately in the case of the generally mouse much like A. insignis but a bit smaller smaller Leyte mouse), but the feet are slightly overall. broader and more robust than in the other The tail scales on the Camiguin mouse are species. All four species have a flat, unpigmented largest, with 12-12.5 scales/cm near the base, nail on the pollex. The relative size and thickness compared to 14-15 scales/cm in A. hylocoetes, follows the same pattern as the hind feet, with A. 13-14 in A. insignis, and 13-15 in the Leyte

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

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 The cranium of Apomys camiguinensis (Figs. 4- same more distally on a given species to about 6) has basioccipital length (28.3-29.0 mm) slight- the midpoint; from there to the tip scale size ly greater than that of A. hylocoetes (28.3- decreases, mostly in the last quarter of the 28.8 mm), clearly more than A. insignis (27.8- length, to about half the length (and one-fourth 28.0 mm) and substantially more than the Leyte the size) of scales near the base of the tail. The Apomys (25.3-26.3 mm); the interorbital width is scales of the tail are dark brown dorsally and proportioned similarly (Table 1). Zygomatic nearly white ventrally on all four species, but the breadth averages greatest in A. hylocoetes transition is most often an abrupt line on the (15.0-15.2 mm), followed by A. camiguinensis Camiguin mouse, often producing a sharply (14.9-15.0 mm), A. insignis (14.7-14.8 mm), and bicolored tail, and most often gradual on the the Leyte Apomys (13.1-13.7 mm), with mastoid other three species. Typically, three hairs grow breadth following the same pattern (Table 1). from beneath each scale, projecting only slightly Nasal length averages substantially greater laterally from the tail; on the basal quarter of the in A. hylocoetes (12.3-12.5 mm) than in A. tail, one of the three hairs is missing from a given insignis (11.6-11.9 mm), A. camiguinensis (11.1- scale about one-third of the time. On A. 11.4 mm), or the Leyte Apomys (10.3-10.7 mm), camiguinensis, the hairs are about one and one- and anterior nasal breadth is similarly patterned third the length of a scale near the base of the (Table 1). Rostral depth averages greatest in A. tail, have about the same length 25% toward the camiguinensis (6.3-6.4 mm), with A. hylocoetes tip, are slightly longer at the midpoint, are about (6.2-6.3 mm) and A. insignis (6.3 mm) slightly one and two-thirds the length of a scale 75% less deep and very similar to each other and the toward the tip, and are about five times the Leyte Apomys generally less deep (5.9-6.1 mm). length of a scale near the tip, where scale Rostral length (Fig. 5) is clearly greatest in A. length has been reduced by about half relative hylocoetes (13.1-13.2 mm), with A. camiguinensis to the base. On A. hylocoetes, hairs are about (11.6-12.0 mm) and A. insignis (11.9-12.0 mm) one and a half the length of a scale near the base similar to each other but much greater than the of the tail, are similar 25% toward the tip, are Leyte Apomys (10.6-11.2 mm). Orbital length about twice the length of a scale near the averages greatest in A. camiguinensis (10.2- midpoint, are about two and a half times 10.3 mm), followed by A. insignis (10.0- the length of a scale 75% toward the tip, and 10.2 mm) and A. hylocoetes (9.7-10.0) and the are more than five times the length of a scale small Leyte Apomys (9.2-9.6). To summarize, A. near the tip. On A. insignis, hairs are one and camiguinensis is characterized by the longest one-third a scale length near the tail's base, skull and deepest rostrum, but A. hylocoetes similar at 25%, one and two-thirds near the has slightly greater zygomatic and mastoid width midpoint, about two and one-half of a scale as well as greater nasal length and breadth and length near 75%, and about three and one-half rostral length. Although the rostrum of A. is in the orbital the length of a scale near the tip. On the Leyte camiguinensis moderate length, mouse, tail hairs are slightly less than the region and braincase are unusually long. The to length of a scale near the base of the tail, similar cranium of A. insignis tends be smaller but at the midpoint and at 75%, and about two times generally similarly proportioned to A. camigui- is smaller in all the length of a scale near the tip. In all four nensis, and the Leyte Apomys it to the species, the hair becomes slightly greater in dimensions, though seems follow pattern A. diameter distally than it is proximally; combined of A. camiguinensis and insignis. with the trend for greater length, this means The maxillary toothrow (Fig. 7) of A. cami- that the tail become more heavily covered with guinensis (6.0 mm) averages nearly identical in and A. hair distally. None of the species shows elongat- length to that of A. hylocoetes (6.0 mm) than ed hairs at the very tip of the tail (i.e., there is insignis (6.0 mm), and all are much greater Palatal no "pencilling"). Hairs on the dorsal surface that of the Leyte Apomys (5.2-5.3 mm). 1 are more heavily pigmented dorsally than breadth at M (Fig. 4) is greatest in A. insignis A. ventrally in all four species. On adults of all four (6.5-6.6 mm), followed by hylocoetes A. species, the dorsal surface of the tip (2-5 mm) of (6.5 mm) and camiguinensis (6.3-6.5 mm) the tail becomes worn, with most scales and and the Leyte Apomys (5.7-5.8 mm). Diastema many hairs absent, leaving a smooth, leathery length (Figs. 4 and 5) in A. camiguinensis surface. (7.3 mm) averages slightly greater than in A.

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE 25 Discussion hvlocoetes (7.2 -7.3 mm), more than A. insignis (7.0 mm), and least in the Leyte Apomys (6.5 of as an 6.8 mm). In other words, the maxillary tooth- The presence Apomys eamiguinensis on a small island, with the rows of the three larger Apomys are all similar, endemic species along additional murine rodent Bullimus (Rick- with the Leyte Apomys having disproportionate- gamay the of A. art et al., 2002) and the Hanging-Parrot (Lor- ly short toothrow, but palate camigui- is that of iculus described in this volume), as noted ncnsis is disproportionately narrow (as sp., indicates the of the Leyte Apomys), and that of A. hvlocoetes is above, clearly importance Island as a center of disproportionately wide. Camiguin unique biological in need of conser- In addition, we note the following qualitative diversity that is worthy and In characters. The incisive foramina (Fig. 5) are vation (Heaney & Tabaranza, 2005). addition, of this confirms widest in A. eamiguinensis and in A. hvlocoetes the distinctiveness species pre- so dictions made on the basis of and longest in A. eamiguinensis and A. insignis, biogeographic that the area of the foramina is greatest in A. models (Heaney, 2004) of the expected presence small on Fur- eamiguinensis. The distance from the posterior of endemic mammals Camiguin. and other edge of the incisive foramina to a line between the ther studies of the mammals, birds, anterior edges of the first maxillary molar is organisms are clearly warranted to determine, difference shortest in A. eamiguinensis, slightly greater in for example, the degree of genetic of which occur on A. insignis, and longest in A. hvlocoetes and from closest relatives (most of the Apomys from Leyte. The braincase of A. Mindanao) as a means of assessing the role eamiguinensis is slightly more elongate and that of colonization and gene flow in determining the Leyte Apomys proportionately most squarish patterns of species richness and endemism in among the four species (Fig. 6). The posterior the Philippines. In other words, Camiguin edge of the bony palate (Fig. 6) extends farthest represents a natural experiment, as a young posterior to the posterior edge of the last oceanic, volcanic island that is near to a large, maxillary molar in A. eamiguinensis, slightly less rich source of species (Mindanao), in which we far in A. hvlocoetes and the Apomys from Leyte. can measure the impact of genetic isolation in and least far in A. insignis. The long axis of the animals and plants of varying vagility under bullae (Fig. 6) is about 45 from the cranial standardized conditions. Such studies are certain midline axis in the Apomys from Leyte, about 40 to produce new insights into the process by c in A. hvlocoetes and A. insignis, and about 35 in which biological diversity is generated in the A. eamiguinensis. The hard palate of A. hvlocoetes Philippines and in other oceanic archipelagos. 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 Acknowledgments anterolabial cusp (probably tl; see Musser & Heaney, 1992, p. 65) that is well developed in A. We thank Rebecca Banasiak for assistance in hvlocoetes, less conspicuous in A. insignis, and photographing skulls, Betty Strack for assistance barely evident in A. eamiguinensis and the Apomys with making scanning electron microscope from Leyte. The first upper molar of A. hvlocoetes images, and Lisa Kanellos for preparation of tends to have a more conspicuous anterolingual figures; Figure 3D was drawn by Jodi Sedlock. cleft than do the other three species (Fig. 7). Both The Protected Areas and Wildlife Bureau of the upper and lower toothrows of A. insignis are the Philippine Department of Environment and most massive (Figs. 7 and 8), with A. eamigui- Natural Resources (denr) provided encourage- nensis somewhat less massive, A. hvlocoetes sub- ment and permits, and the Region 10 office of stantially less so, and the Apomys from Leyte the denr provided logistical support. Specimens smallest overall. All four species have the canal for comparison were kindly loaned by G. Hess for the infraorbital branch of the stapedial artery (dmnh), P. Myers (ummz), and M. Carleton, L. partially open and part of the artery exposed on Gordon, and H. Kafka (usnm). The manuscript the ventral surface of each pterygoid plate (Fig. 6), was much improved by comments from D. S. as noted Musser by (1982). Balete, G. G. Musser, P. Myers, and E. A. Ecology Sec Heaney et al. (2006) for ecolog- Rickart. Funding was provided by the Marshall ical information Field Fund, Ellen Thorne Smith Fund, and the

26 FIELDIANA: ZOOLOGY Barbara Brown Fund for Mammal Research of Heaney, L. R., and B. R. Tabaranza, Jr. 1997. A preliminary report on mammalian and the Field Museum of Natural History. diversity conservation status of Camiguin Island, Philippines. Sylvatrop, 5(1995): 57-64.

. 2006. Introduction to the mammal and land bird faunas of Camiguin Island, Philippines, pp. 1- Literature Cited 13. In Heaney, L. R., ed., The Mammals and Birds of Camiguin Island, Philippines, a Distinctive Center of J. C. 1971. The of 1. Brown, description mammals, Biodiversity. Fieldiana Zoology, n.s., 106: 1-72. The external characters of the head. Mammal Heaney, L. R., B. R. Tabaranza, Jr., D. S. Balete, Review, 1: 151-168. and N. Rigertas. 2006. Synopsis and biogeography Brown, J. C, and D. W. Yalden. 1973. The of the mammals of Camiguin Island, Philippines, description of mammals, 2. Limbs and locomotion pp. 00-00. In Heaney, R. L., ed., The Mammals and of terrestrial mammals. Mammal Review, 3: Birds of Camiguin Island, Philippines, a Distinctive 107-134. Center of Biodiversity. Fieldiana Zoology, n.s., 106: 28-48. Goodman, S. M., and N. R. Ingle. 1993. Sibuyan Island in the Philippines—Threatened and in need of Mearns, E. A. 1905. Descriptions of new genera and conservation. Oryx, 27: 174-180. species of mammals from the Philippine Islands. of the United States National Museum Heaney, L. R. 1984. Mammals from Camiguin Island, Proceedings of Natural History, 28: 425-460. Philippines. Proceedings of the Biological Society of Washington, 97: 119-125. Musser, G. G. 1982. Results of the Archbold Expeditions, No. 108. The definition of Apomys, . 1986. Biogeography of mammals in Southeast a native rat of the Philippine Islands. American Asia: Estimates of rates of colonization, extinction, Museum Novitates, 2746: 1-43. and speciation. Biological Journal of the Linnean G. and L. R. Heaney. 1992. Society, 28: 127-165. Musser, G., Philippine rodents: Definitions of Tarsomys and Limnomys 1991. An analysis of patterns of distribution plus a preliminary assessment of phylogenetic and species richness among Philippine fruit bats patterns among native Philippine murines (Murinae, (Pteropodidae). Bulletin of the American Museum Muridae). Bulletin of the American Museum of of Natural History, 206: 145-167. Natural History, 211: 1-138. -. 1993. Biodiversity patterns and the conserva- Rickart, E. A., and L. R. Heaney. 2002. Further tion of mammals in the Philippines. Asia Life studies on the chromosomes of Philippine rodents Sciences, 2: 261-274. (Muridae: Murinae). Proceedings of the Biological 2004. Conservation biogeography in oceanic Society of Washington, 115: 473^87. 345-360. In Lomolino, M. V. and archipelagoes, pp. Rickart, E. A., L. R. Heaney, P. D. Heideman, and of L. R. Heaney, eds., Frontiers Biogeography: New R. C. B. Utzurrum. 1993. The distribution and Directions in the of Nature. Sinauer Geography ecology of mammals on Leyte, Biliran, and Maripipi Associates, Sunderland. Islands, Philippines. Fieldiana Zoology, n.s., 72: Heaney, L. R., D. S. Balete, L. Dolar, A. C. 1-62. P. C. N. R. M. Alcala, A. Dans, Gonzales, Ingle, Rickart, E. A., L. R. Heaney, and B. R. Tabaranza, Lepiten, P. S. Ong, W. L. R. Oliver, E. A. Rickart, Jr. 2002. Review of Bullimus (Muridae: Murinae) B. R. and R. C. B. Utzurrum. Tabaranza, Jr., and description of a new species from Camiguin 1998. of the fauna of the A synopsis mammalian Island, Philippines. Journal of Mammalogy, 83: Philippine Islands. Fieldiana Zoology, n.s., 88: 1- 421^136. 61. Ruedas, L. A. 1995. Description of a new large-bodied Heaney, L. R., and J. C. Regalado, Jr. 1998. species of Apomys Mearns 1905 (Mammalia: Vanishing Treasures of the Philippine Rain Forest. Rodentia: Muridae) from Mindoro Island, Philip- The Field Museum, Chicago. pines. Proceedings of the Biological Society of 108: 302-318. Heaney, L. R., and E. A. Rickart. 1990. Correlations Washington, of clades and clines: Geographic, elevational, and Steppan, S., C. Zawadski, and L. R. Heaney. 2003. A phylogenetic distribution patterns among Philippine molecular assessment of phylogenetic relationships mammals, pp. 321-332. In Peters, G. and R. and patterns of phylogenesis in the Philippine murid Hutterer, eds., Vertebrates in the Tropics. Museum rodent Apomys. Biological Journal of the Linnean Alexander Koenig, Bonn. Society, 80: 699-715.

HEANEY AND TABARANZA: A NEW SPECIES OF FOREST MOUSE 27 of Synopsis and Biogeography of the Mammals Camiguin Island, Philippines

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

Abstract

Biodiversity surveys in the 1960s and 1990s on Camiguin Island, a geologically young, the volcanically active oceanic island surrounded by deep water, have demonstrated presence native of 24 species of land mammals. Three species (one insectivore and two rodents) are not 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 Tne Islands Department of Biology, Iligan Institute of Tech- Philippine 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 2 from less than one to Laksambuhay Conservation. Inc., 10241 Mt. hectare over 100,000 km , Bulusan Street, U-2. Los Banos, with Laguna. Philippines. geological age varying from under 1 million

28 FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, PP. 28-48 to over 40 million These years years. islands indicated in a brief preliminary report (Heaney & sets of represent many historically distinct geo- Tabaranza, 1997), we found eight additional units of logical remarkably varied origins; some species on the island that are widespread in the had connections land-bridge to the Asian main- Philippines, plus two previously unknown en- land in the of past (those the Palawan group), demic species of rodents. The purpose of this but most are purely oceanic in origin (Heaney, paper is to summarize the results of the 1994 and 1986, 1991b, 2000; Heaney & Rickart, 1990; 1995 mammal surveys and integrate those data Hall, 1998, The mammals that 2002). have with information from the 1960s, including evolved in this diverse archipelago include at information on habitat associations, relative least 1 1 1 endemic to the out species archipelago abundance, and ecology of the species. 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 Methods the endemic species occur on the large islands of Luzon, Mindanao, Mindoro, Negros, and Pala- Prior Reports wan (e.g., Heaney, 1986, 1993, 2000; Heaney et al., 1998; Rickart et al., 1998), significant The first report of mammals from Camiguin numbers occur on the smaller islands as well, Island was that of Gray (1843), who reported Paradoxurus especially those surrounded by deep water (e.g., hermaphroditus. Three field teams Heaney, 1986, 2004; Goodman & Ingle, 1993; from Silliman University led by Dioscoro S. Heaney & Tabaranza, 1997; Musser et al., 1998). Rabor collected mammals on Camiguin in 1967, As noted by Heaney and Tabaranza (2006a), 1968, and 1969; specimens were deposited at the 2 Camiguin, an island of 265 km located about Delaware Museum of Natural History (dmnh) 10 km north of Mindanao in the Bohol Sea, is and Royal Ontario Museum (rom); for details, one such deep-water island, with a minimum see Heaney and Tabaranza (2006a). Several depth to Mindanao of 385 m. It is steeply specimens were reported by Peterson and Fenton mountainous, with several active volcanic cones (1970); all known specimens from the 1960s were that reach to a maximum elevation of about examined and summarized by Heaney (1984). All 1600 m. A series of biological surveys on data included in this paper from the 1960s Camiguin in the late 1960s that focused on birds specimens are based on data in Heaney (1984), as (see Balete et al., 2006) also yielded some except noted below. mammal specimens, and an earlier report on those surveys (Heaney, 1984) concluded that the Recent Data island had no endemic mammal species and was depauperate. Subsequent studies on other islands Field studies were conducted during three made us suspect that those earlier surveys were periods in 1992, 1994, and 1995; general methods incomplete because so few mammal species had and site descriptions are given in Heaney and been obtained and because the number of Tabaranza (2006a). Sampling during 1995 fol- nonvolant mammal specimens was small (thus lowed methods used on Leyte, Luzon, Negros, indicating limited sampling effort). Further, on and other islands (Heaney et al., 1989, 1999; the basis of biogeographic patterns elsewhere in Rickart et al., 1991, 1993) to facilitate quantita- the Philippines, we predicted the presence of tive comparisons. Nonvolant small mammals several endemic small mammals on Camiguin were caught in traps; during 1995, all traps were (Heaney, 2004). To investigate the hypotheses Victor rat snap traps. Most were baited with that the previously measured species richness was fresh fried coconut coated with peanut butter, low because of incomplete surveys and that but a few were baited with live earthworms. about two endemic species should be present, we During 1994, several National live traps were returned to Camiguin briefly in 1992 and more used, in addition to Victor rat traps, and were extensively in 1994 and 1995 to conduct addi- baited with coconut bait. Bats were captured in tional mammal inventories in all the major 12-m mist nets. Voucher specimens were pre- habitats along the elevational gradient, especially pared in fluid or as skeletons and have been by trapping small mammals at higher elevations deposited at The Field Museum of Natural where there were few records from the 1 960s. As History (fmnh), National Museum of the Phi-

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND 29 forest State elevation (Site 5), and in primary montane lippines (nmp), and Mindanao University- Most at 1275 m (Site 7). It was most often trapped lligan Institute ol' Technology (msu-iit). in- under tree roots and live vegetation. None were specimens were autopsied for reproductive taken in areas at 150 m (Site 3), or in formation. The si/e of embryos was measured as agricultural animals forest at 1475 m (Site 8, where there was crown to rump length (CRL). Subadult mossy Table 1 This use of habitat is are defined here as those that have not completed limited sampling; ). unfused consistent with data from islands on Greater cranial growth, especially those having et et al., basicranial sutures; these young animals have Mindanao (Rickart al., 1993; Heaney than that pelage that is usually softer and grayer unpubl. data). three adult females were of adults and are noticeably lower in weight and In 1994 and 1995, was with a females are usually nulliparous. Young adults trapped; one pregnant single embryo = 10 are older; they have nearly completed cranial (CRL mm). A multiparous, nonpregnant 13 an adult male 7 growth but have not yet reached adult weight female weighed g; weighed g. or are cranial and external measurements and, usually, have not yet reproduced Both are within stated for Mindanao pregnant for the first time. Adults have complet- (Table 2) ranges smaller ed cranial growth and adult pelage, and usually (Heaney & Ruedi, 1994) but are slightly and the females are multiparous. Comments on than those of series taken on Biliran, Leyte, distribution and use of scientific names are based Maripipi (Rickart et al.,—1993). on Heaney et al. (1998) unless additional sources Specimens Examined Total 5. Site 4 (2 fmnh, are mentioned. Records of specimens examined 1 msu-iit); Site 5 (1 fmnh); Site 7 (1 fmnh). are summarized at the end of each account; such Suncus murium 1 766) summaries include site number and the number (Linnaeus, The Asian house-shrew occurs widely in Asia of specimens (in parentheses). and Indo-Australia; it now occurs throughout External measurements and weights reported the it is not native to the here were taken in the field by members of Philippines, though It is abundant in urban and the field team on fresh animals. Cranial mea- country. agricultural areas; on islands with low mammal species surements were taken by Heaney with digital richness such as Negros, it is sometimes abun- calipers graduated to 0.01 mm. Comparisons of dant in disturbed forest and occasionally in cranial measurements are to published records of forest et al., 1987; Heaney et specimens measured in the same manner by primary (Heideman al., 1989, 1991), but on islands of average species Heaney. richness, it is usually rare or absent from forest (Heaney et al., 1989, 1998. unpubl. data; Rickart et al., 1993). Accounts of Species A single subadult specimen from Mt. Tim- poong Peak was available previously (Heaney Order Insectivora 1984). In 1994-1995, we captured this species Family Soricidae—Shrews from 150 to 1475 m, and it was the most Crocidura beatus Miller, 1910 common species at the three highest sites, all in The Mindanao shrew is widespread on islands primary forest (Table 1, Fig. 1). It was especially in the Mindanao Faunal Region (Heaney & abundant in montane forest at 1275 m (Site 7). It Ruedi, 1994; Heaney et al., 1998); these are the was moderately abundant in primary mossy first records from an island that was not part of forest (Site 6; elev. 1300 m) and in lower mossy the late Pleistocene island of Greater Mindanao forest at 1475 m elevation (Site 8) but was much (Heaney. 1986). This shrew has most often been less common in heavily disturbed lowland found in primary forest, especially at higher agricultural land at 150 m (Site 3). This pattern elevations; is usually uncommon in secondary of abundance is quite different from that on the forest; and is absent outside of forest (Heaney et species-rich islands of Biliran, Leyte, Luzon, al.. 1989; Rickart et al., 1993). Maripipi, and Mindanao, where specimens were Crocidura beatus was trapped on Camiguin at never caught in primary forest (Heaney et al., three forest sites in May 1994 and March 1995 1989, 1999, unpubl. data; Rickart et al., 1993) (Fig. 1. Table 1). It was uncommon in secondary but similar to the species-poor island of Negros, lowland forest at 1000 m elevation (Site 4), in where S. murinus was abundant in transitional disturbed lower montane forest at 1200 m mossy/montane forest and in mossy forest at

30 FIELDIANA: ZOOLOGY *=1960s 1800 i- =1990s

1600 =" maximum elevation — 1400 mossy primary J 1 1 1. 1 1 ~ 1200 " montane disturbed O 1000 — L re _ lowland | 800

600 — no forest

400 —

200

W CO c/> 11 o .c II c t c 5 c II •a 5) £ O 03 o cc 5 ^ Q..C II 0) c OQ "3 o .5)

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 Specimens Examined Total 78. Site 3 (2 msu-

Sites 6 and 7. iit); Site 6(10 msu-iit); Site 7 (56 fmnh); Site 8 (9 Suncus murinus was most often trapped in fmnh); Site 15 (1 dmnh). runways or clear areas beneath fallen and rotting logs, under roots of trees, or under horizontal Order Chiroptera trunks of live trees as well as in runways near Family Pteropodidae—Fruit Bats large boulders. Many were caught during day- Cynopterus brachyotis (Muller, 1838) light hours. The common short-nosed fruit bat is wide- Five adult females with a mean weight of 32 ± spread in Southeast Asia and is common = 4.5 g (range 27-39 g) were pregnant; litter throughout the Philippines. It ranges from sea sizes for four of these were one, two, three, and level to at least 1250 m and is typically found in three. Thirteen nonpregnant parous females agricultural areas; it is also common in second- (those with large mammae) weighed an average ary lowland forest but usually rare in primary = et of 28.8 ± 4.1 g (range 22-35 g), and nullipa- forest (Heaney al., 1998). rous females (those with small mammae) Our limited netting on Camiguin during 1994- = = weighed 20.6 ± 2.2 g (range 17.5-23 g, N 1995 (Table 3) indicated that C. brachyotis was 10). Adult males (defined as those with large abundant in a highly disturbed lowland agricul- testes) had a mean weight of 36 ± 5.8 g (range tural area at 10 m elevation (Site 1) and was = in lowland 24-48 g, N 27). Males are conspicuously common a heavily disturbed agricul- larger than females in this species (Table 2). tural area at 100 m (Site 2; Table 3). It was less

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND 31 in in disturbed lowland agricultural Tabu 1. Numbers of nonvolant small mammals captured traps heavily disturbed lower montane forest (Site 5), primary mossy forest (Site area (Site 3) secondary lowland forest (Site 4). forest on Island. The numbers of captures 6) primary montane forest (Site 7), and lower mossy (Site 8) Camiguin See and Tabaranza (2005a) for full site descriptions. Asterisks per KM) trap-nights are given in parentheses. Heaney see Methods. mark species presumed to be present; r<

e a Ctf 00 60 £ s .5 >

SI es

C-o

T3 .;; ooa •He o o

oo on ±j P t3 a " c O oo 2 Sc5* E

5.SE

S o.E

Q*2 a +i g § — 6 E c a B

^Z 1 E

HIS area Tabu: 3. Numbers of fruit bats captured in mist nets in a lowland agricultural (Site 1), heavily disturbed disturbed lower montane forest and lowland agricultural area (Site 3), secondary lowland forest (Site 4), 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).

Site I, Site 3, Site 4, Sites 5 and 6, Site 7, Scientific name 10 m 150 m 1000 in 1200-1400 m 1275 m

( 'ynopterus brach vol is a'S

C/3 of fruit bats, in cranial and external measurements is slight, endemic species Philippine pygmy and Camiguin specimens tend to cluster consistently Haplonycteris fischeri Otopteropus eartilago- where the was first detected within the upper ranges of most features nodus, phenomenon Heideman et al., 1993; Heide- measured (Table 4; Heaney and Rabor, 1982; (Heideman, 1989; man Powell, 1998). Additionally, this condi- Heaney et al., 1991, 1999, unpubl. data; Rickart & exhibited et al., 1993). tion was apparently by primiparous females them to Specimens Examined—Total 56. Site 1 (15 young adult only, allowing give in their first which had the 1 Site 4 Site 5 birth once year, fmnh); Site 3 ( msu-iit); (4 msu-iit); only of them to (8 msu-iit); Site 6 (1 msu-iit); Site 11(12 dmnh); effect enabling synchronize breeding the Site 13 (5 dmnii); Site 14 (2 dmnii); Site 17 (8 with the adult females following year (Heide- dmnii). man & Powell, 1998). Males are somewhat larger than females in Peters, Ptenochirus jagori ( 1861) most cranial and external dimensions, as on The fruit bat is a common Philippine musky Biliran, Leyte, and Maripipi (Table 4; Rickart et endemic, the archipelago occurring throughout al., 1993). Cranial and external measurements with the of the and exception Batanes/Babuyan (Table 4) are noticeably larger than those for Palawan faunal from sea level to at least regions series from Catanduanes and southern Luzon 1800 m et al., 1998). Our limited netting (Heaney and were similar to those from Biliran, Leyte, on Camiguin showed Ptenochirus jagori to be and Maripipi (Heaney, 1984; Heaney et al., 1991, common in a lowland agricultural area at 10 m 1999; Rickart et al., 1993). elevation (Site 1), less common in disturbed Specimens Examined—Total 46. Site 1 (8 lowland forest at 1000 m (Site 4), and scarce in fmnh); Site 4(16 msu-iit); Site 6 (1 msu-iit); Site primary mossy forest at 1300 m (Site 6; Table 3). 11 (6 dmnh); Site 13 (6 dmnh); Site 17 (9 rom). Combined with records from the 1960s, these data indicate that the species is widespread in Pteropus hypomelanus Mearns, 1905 lowland and montane rain forest from sea level The common island flying fox occurs from to at least 1200 m (Fig. 2), though probably its Thailand to Australia and is found throughout declines with elevation and abundance increasing the Philippines with the exception of the Palawan with levels of disturbance as increasing (Table 3), faunal region. It is often common in agricultural noted elsewhere (Heaney et al., 1989, 1999, areas from sea level to ca. 900 m and is absent in Rick- unpubl. data; Heideman & Heaney, 1989; primary forest (Heideman & Heaney, 1989; art et 1993; al., Lepiten, 1997). Heaney et al., 1991, 1998; Utzurrum, 1992; Three adult netted in 1992 and females, May Rickart et al., 1993). Records from the 1960s 1994, an of 70 = 68- weighing average g (range document it from Camiguin at elevations from 75 were with a each g), pregnant single embryo about 250 to 1500 m (Fig. 2), but we netted none (CRL = 5 10 Three females mm). nonpregnant in the 1990s; because this species typically flies with mammae had an of large average weight above the canopy and our nets were set not more 73.5 = 72-74 and two g (range g), nulliparous than about 4 m above the ground, our failure to females 67 and 68.5 Eleven adult weighed g. catch any does not necessarily indicate any males had a 73.2 ± 4.7 mean weight of g (range in their abundance. = = change 64-78 g, N 11). Pregnant females of P. External and cranial measurements show only jagori have been recorded also in May on Luzon slight variations with those of specimens from (Mt. Isarog) (Heaney et al., 1999). On Mindanao Dinagat, Panay, Leyte, and Maripipi (Heaney & (Kitanglad Range), pregnant females were re- Rabor, 1982; Rickart et al., 1993). corded in March, May, July, and August and Specimens Examined—Total 8. Site 10 (2 lactating females in May to June and August to dmnh); Site 12 (1 dmnh); Site 13 (2 dmnh); Site December (Heaney et al.. unpubl. data). Heide- 17 (3 ROM). man and Powell (1998) found that on Negros Island, P. jagori gives birth to a single young Pteropus pwnilus Miller, 1910 twice each the first in late or year: March early The little golden-mantled flying fox is endemic and the second in It was further to the April August. Philippines (aside from a single population discovered that this species undergoes delayed on Miangas Island. Indonesia, adjacent to and that lasts implantation early development Mindanao) and occurs throughout the archipel- for five shorter than in two months, other ago, with the exception of the Batanes/Babuyan

36 FIELDIANA: ZOOLOGY and Palawan regions (Heaney et al., 1998). is most often encountered from lowlands to at from Previously reported Camiguin as P. tablasi, least 1350 m in agricultural lands to primary the revised species was to include P. tablasi and lowland and montane forest and occasionally P. balutus as under synonyms Pteropus pumilus roosts in caves (Heaney et al., 1991, 1999; (Klingener & Creighton, 1984). It is associated Rickart et al., 1993). Two groups that differ in with primary and good secondary lowland forest size and habitat are recognized within this from sea level to about 1100 m, and it is species: a smaller morphotype, designated R. uncommon outside of forest. Additionally, it is arcuatus—s, that occurs in the lowlands or most common on small islands and is uncommon disturbed habitats, and a larger one, designated to rare on islands. — larger Pteropus pumilus often R. arcuatus 1, found in forest at higher eleva- is netted in clearings or on ridgetops (Heideman tions (Ingle & Heaney, 1992). & Heaney, 1989; Utzurrum, 1992; Rickart et al., In May 1994, we netted this species at 1000 m 1993). elevation in disturbed lowland forest (Site 4). Of We did not encounter P. pumilus during the two adult females netted, one (15 g) was = 1990s, but the 1967-1969 surveys obtained 42 pregnant with one large embryo (CRL individuals at four sites from ca. 250 to nearly 28 mm), and the other was lactating (12.5 g). 1000 m elevation (Fig. 2). As with Pteropus Pregnancies in this species were recorded also in hypomelanus, this species usually flies above the March on Luzon (Mt. Isarog) and in April on canopy, so our failure to catch any in the 1990s Biliran, Leyte, and Maripipi (Rickart et al., 1993; does not necessarily indicate a change in their Heaney et al., 1999). status on the island. Cranial and external measurements (Table 5) Specimens Examined—Total 42. Site 11 (19 of the Camiguin specimens are consistently in dmnh); Site 13 (2 dmnh); Site 14 (1 dmnh); Site larger than those series from Biliran, Leyte, 17 (20 ROM). Luzon (Mt. Makiling), Maripipi, and Mindanao (Mt. Kitanglad), which all fall within the Emballonuridae—Sheath-tailed Bats Family dimensions of the smaller morphotype, R. Emballanura alecto and (Eydoux Gervais, 1836) arcuatus—s (Ingle & Heaney, 1992; Rickart et The sheath-tailed bat is a common Philippine al., 1993; Heaney et al., 1999, unpubl. data). cave-dwelling species that occurs throughout Instead, their external and cranial dimensions are most of the and is also known from Philippines comparable to the larger morphotype, R. arcua- the and Sulawesi Borneo, Moluccas, (Koopman, tus—1 (Ingle & Heaney, 1992). Systematics in this It has been recorded in lowland areas 1989). only "species" are badly in need of detailed study. and in disturbed forest (450 m below) and Specimens Examined—Total 10. Site 4 (4 msu- areas with scattered remnant agricultural forest, iit); Site 13 (6 dmnh). with most captures recorded from caves, under K. 1905 large boulders, or in man-made tunnels (Heaney Rhinolophus inops Anderson, The forest horseshoe bat is common et al., 1991, 1999; Rickart et al., 1993). We did Philippine to abundant in lowland and montane not record this species during our fieldwork in primary forest from sea level to 2250 and is 1994 and 1995, but in 1967 specimens were taken m usually rare in forest and forest from Tag-ibo Cave at 400 ft (ca. 120 m) and at secondary mossy et al., 1991, 1998, 1999; Rickart et al., 1400-3300 ft (ca. 400-1000 m) on Mt. Mamba- (Heaney in our of jao (see also Heaney, 1984). 1993). Improvements understanding lead us to the Comparison of external and cranial measure- Philippine Rhinolophus reidentify from Site as R. ments with series from Leyte and Biliran shows single specimen 13, reported (1984), as R. inops. We little variation (Table 4; Rickart et al., 1993). subrufus by Heaney netted two additional males in disturbed lowland Specimens Examined—Total 4. Site 17 (2 forest at 1000 m (Site 4). Cranial and external rom); Site 18 (2 rom). measurements (Table 5) are slightly larger in Family Rhinolophidae—Horseshoe-nosed Bats most dimensions than those of the series from Rhinolophus arcuatus Peters, 1871 Biliran, Leyte (Rickart et al., 1993), and Mind- The arcuate horseshoe bat is widespread from anao (Mt. Kitanglad; Heaney et al., unpubl. Sumatra to New Guinea and throughout the data); we suspect that regional morphs corre- Philippines (Heaney et al., 1998), including the sponding to the Pleistocene islands (Heaney, Palawan faunal region (Esselstyn et al., 2004). It 1986) are present.

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND 37 E o , O N — o S E"

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U o Specimens Examined—Total 3. Site 7 (2 there was limited sampling at all of these fmnh); Site 13 (1 dmnh). (Table 1). Most specimens were taken from traps placed in primary montane forest underneath Order Primates root tangles and beneath fallen and rotten logs Family Cercopithecidae—Monkeys on a moderate slope. All individuals were Macaca fascicularis (Raffles, 1821) captured at night. The long-tailed macaque occurs from Burma A pregnant female taken on 29 May 1994 = to Timor and is found throughout the Philip- weighed 40.5 g and had a single embryo (CRL pines (Fooden, 1995; Heaney et al., 1998). It is 30 mm). Adult females with large mammae but = found in agricultural areas near forest, in second- not pregnant weighed 39 ± 2.1 g (range 37.5- = growth and secondary forest, as well as primary 42 g, N 5); two of these, taken 23 May 1994 forest from sea level to at least 1800 m in and 17 March 1995, each had two placental lowland and montane forest (Goodman & Ingle, scars. Nulliparous females (with small mammae) 1993; Rickart et al., 1993; Heaney et al., 1999). were lighter and had an average weight of 35 g = We did not collect any specimens, but in March (range 34-36). Females have two pair of 1995 we obtained the skull of an adult individual inguinal mammae. These data indicate a litter from a hunter who had killed the macaque in size between one and two. Males with scrotal a forested area above Mahinog a few years testes had a mean weight of 41.3 ± 4.7 g (range = = earlier. Also in March 1995, a local hunter who 36.5^48 g, N 11); testis size ranged from 15 was a member of our team saw two macaques X 16 mm to 8 X 12 mm. Males with abdominal at = a short distance from our camp 1100m testes weighed 33.0 ± 5.0 g (range 28-34 g, N = elevation (Site 7) in primary montane forest. 5) and had testis size of 5 X 8 mm. He reported that the macaques were fairly Specimens Examined—Total 25. Site 4 (14 in of common what remained lowland primary msu-iit), Site 6 (2 msu-iit); Site 7 (9 fmnh). forest and good secondary forest at that time and Bullimus Rickart, and Tabaranza, were actively hunted. gamay Heaney 2002 Specimens Examined—Total 1. Above Mahi- A unknown of Bullimus, nog (1 fmnh). previously species recently described as B. gamay (Rickart et al., Order Rodentia 2002), was captured on Camiguin in May 1994 and March 1995. The forest rat was Family Muridae—Rats and Mice Camiguin to and Apomys camiguinensis Heaney and Tabaranza, incorrectly assigned Tarsomys by Heaney 2006 Tabaranza (1997) and was designated "Bullimus A" et al. (1998). It was trapped A previously unknown species of Apomys sp. by Heaney from 900 to 1475 m 1, Table but not in (Musser, 1982) was trapped on Camiguin in (Fig. 1) disturbed lowland land from May 1994 and March 1995 (Heaney & Tabar- heavily agricultural sea level to 300 m Most were anza, 1997) and described in this volume (Site 3). specimens taken from in beneath root (Heaney & Tabaranza, 2006b). The Camiguin traps placed runways and rotten or near rocks. This forest mouse is moderately large for the genus, tangles logs large was most common in primary montane averaging about 40 g. It is marked by a number species forest at 1275 m 7). It was less common in of subtle morphological characters, and molec- (Site disturbed lower montane forest at 1000-1300 m ular data show it to be most closely related to and in forest at 1200- Apomys hylocoetes and A. insignis from Mind- (Site 5) primary mossy 1400 m It was rare in lowland anao and to an unnamed species from Leyte and (Site 6). secondary forest at 1000 m (Site 4) and in lower mossy Biliran (Steppan et al., 2003; Heaney & Tabar- forest at 1475 m Bullimus anza, 2005b). This species was common (Table 1, (Site 8). bagobus occurs in similar habitat on Mindanao but also Fig. 1) in disturbed lowland forest at 1000 m occurs at lower elevation where forest (Site 4) and in primary montane forest at 1275 m good et (Site 7) and relatively uncommon in primary persists (Heaney, 2001; Heaney al., unpubl. this that B. should also be mossy forest at 1200-1400 m (Site 6). It was data); suggest gamay at lower elevations on not found in heavily disturbed lowland agri- sought Camiguin. females with mammae cultural land (Site 3, elev. 150 m), lower Parous large weighed 370 = 360-390 N = montane forest (Site 4, elev. 1000-1300 m), or an average of g (range g, 3), while adult females with small lower mossy forest (Site 8, elev. 1475 m), but nulliparous young

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND 39 Cranial and external measurements mammae were lighter, having a mean weight of (Table 6) = = are similar to but than those from 276 ± 53.7 g (range 210-345 g, N 5). Adult slightly larger males with scrotal testes had an average weight Leyte and Mindanao (Kitanglad Range and Mt. = - Rickart et al., 1998; et al., of 402 g (range 240-500 g; N 3) and had Apo; 1993, Heaney testis size from 21 x 30 mm to 27 X data). ranging unpubl. — 35 mm). Among the nine individuals taken in Specimens Examined Total 5. Site 4 (2 msu- May 1994, the smallest was a nulliparous young iit); Site 5 (1 msu-iit); Site 7 (2 fmnh). female weighing 210 g, while among the 11 individuals captured in March 1995, two were Rattus everetti (Gunther, 1879) juveniles, a male weighing 120 g and a female The common Philippine forest rat is a wide- weighing 125 g. No pregnant females were taken spread endemic species (excluding the Babuyan/ in May 1994 and March 1995, but a lactating Batanes, Palawan, and Sulu groups) found in female (360 g) was taken on 18 March 1995, and primary and disturbed lowland, montane, and a female taken on 29 May 1994 had three mossy forest from sea level to 2400 m (Heaney et placental scars. This suggests that females give al., 1991, 1998, 1999; Musser & Heaney, 1992; birth in February and March. Three females had Rickart et al., 1993). three pairs of mammae and one had four pairs, During 1994-1995, we found this 250-350 g indicating that litter size most likely does not rodent to be fairly common at all sampling sites exceed three. from disturbed lowland forest at 1000 m, Specimens Examined—Total 20. Site 4 (4 msu- through montane forest to mossy forest at iit); Site 5 (2 msu-iit); Site 6 (3 msu-iit); Site 7 (10 1475 m; specimens from the 1960s were taken emnh); Site 8 (1 fmnh). at 500 m (Table 1, Fig. 1); on Mindanao, it occurs down to sea level (Heaney, 2001; Heaney Crunomys melanius Thomas, 1907 et al., unpubl. data). It was trapped under root The Mindanao shrew-mouse was previously tangles and beneath fallen logs, in hollow spaces known from and Mindanao from only Leyte beneath live trees, and near mounds of soil and near sea level to 1550 m, in apparently primary rotting leaves. lowland rain it is rare in collections forest; Five parous adult females from May 1994 and (Musser & 1992; et al., 1998; Heaney, Heaney March 1995 had a mean weight of 271 ± 28.8 unpubl. data; Rickart et al., 1998). On Camiguin, = = (range 240-300 g, N 5). During the same we found it to be uncommon in disturbed heavily months, five nulliparous females (with adult lowland land at 1000 m in agricultural (Site 4), pelage) weighed 188 ± 21.4 g (range = 160- disturbed lower montane forest at 1200 m (Site 215 g), and four females with juvenile pelage 5), and in primary montane forest at 1275 m = weighed 116 ± 31.0 g (range 77-150 g). (Site 7; Table 1, 1). It was beneath Fig. trapped Females have four pairs of mammae, indicating rotten and wood in an area with few logs tangles moderate litter size. None were pregnant, and we dead leaves and little moss. It occurs down to sea were unable to count placental scars; previous level in forest on Mindanao et good (Heaney al., studies also produced little data on litter size 1998; Heaney, 2001) and should also be sought (Heaney et al., 1999, unpubl. data). Males with at lower elevations on Camiguin. scrotal testes, recorded in May 1994 and March Two of the five we were specimens captured 1995, had a mean weight of 288 ± 69.8 g (range females. An adult female taken on 17 March = = 210 375 g, N 4), and the largest males had 1995, with enlarged mammae and weighing 71 g, testes size of 33 X 52 mm and 30 X 50 mm. had birth; it had three recently given placental Three males with abdominal testes, during the scars (one left, two and the uterus was right), same months, weighed an average of 207 g swollen. The other female = (62.5 g), probably (range 160-295 g). nulliparous, had small mammae. Two adult Cranial and external measurements (Table 6) males (60 g and 72 g) had scrotal testes X are (15 comparable to those from Biliran, Leyte, and 8 mm); the third male, taken on 24 May 1994, Maripipi but slightly smaller than series from was a small (28 with abdominal testes. juvenile g) Dinagat, Mindanao (Kitanglad Range), and These scanty data imply that March, and April, Siargao (Heaney & Rabor, 1982; Heaney et al., May are months of Fe- reproductive activity. 1991, 1999, unpubl.; Rickart et al., 1993). males have two pairs of mammae, that indicating Specimens Examined—Total 28. Site 4 (6 msu- litter size is small. Site 5 iit); (2 msu-iit); Site 6 (3 msu-iit); Site 7 (8

40 FIELDIANA: ZOOLOGY -*- ^ O ON

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2 89 et On we 1 Site 12 Rickart al., 1993). fmnh); Site 8 (2 fmnh); Site 1 (4 dmnii); data; Camiguin, found R. tanezwni to be abundant in (1 dmnh); Site 13 (2 dmnii). very heavily disturbed agricultural land at 150 m (Site 3; Rutins exulans (Peale, 1848) Table 1). It was less common in secondary Known as the Polynesian rat or the spiny lowland forest at 1000 m and in disturbed lower is to the rice-field rat, R. exulans not native montane forest at 1000 m (Site 4) and 1200 m accidental Philippines; through mostly dispersal (Site 5), and a single individual was trapped in it from to Easter by humans, occurs Bangladesh primary mossy forest at 1400 m (Site 6). Four Island and throughout the Philippines, especially nestling juveniles were found in a new slash-and- as a pest in agricultural areas. On Camiguin, we burn clearing near Site 7, but no R. tanezwni found this small (60-75 g) rat to be moderately were caught in adjacent mature forest. Taken common in heavily disturbed lowland agricul- together with records from the 1960s, these data tural land at ca. 150 m (Site 3) and in primary indicate that R. tanezwni occurs from sea level to montane forest at 1275 m (Site 7) but not at the highest peaks on Camiguin, although it tends other sites. Records from the 1960s also show it to be most common at lower elevations and in to be present in mossy forest at ca. 1500 m. This disturbed habitats. distribution is similar to that found on Negros, Pregnant females had a mean weight of 126 ± = = where it occurs in high-density agricultural areas 57.6 g (range 63-200 g, N 4). Females with and in mossy forest at an elevation of 1500- large mammae but not pregnant had a mean = = 1650 m (Heideman et al., 1987; Heaney et al., weight of 151 ± 16.7 g (range 130-169 g, N 1989). Both Camiguin and Negros have very few 9); three of these had placental scars (three, four, native rodents, and this may influence the ability and eight scars). Nulliparous females had a mean = of nonnative species to invade the forest since the weight of 93 ± 39.8 g (range 67-160 g). Males nonnatives are absent or very rare in mature with scrotal testes had a mean weight of 169.4 ± = = forest on species-rich islands such as Leyte, 22.7 g (range 141-199 g, N 8); a subadult Luzon, and Mindanao (Heaney et al., 1989, with scrotal testes weighed 105 g. These data 1999. unpubl. data; Rickart et al., 1993). indicate large litter size (up to eight) and very In March 1995, one nulliparous female weigh- early reproduction, as is typical for the species ing 41 g was trapped. Two adult males (71-76 g) (Barbehenn et al., 1973). External and cranial with scrotal testes and two young adults (49- measurements (Table 6) are comparable to those 61 g) with scrotal testes were trapped at the same of Biliran, Leyte, Maripipi, and Mindanao, with time. They typically become reproductive at only slight variation. — a young age and have many large litters each Specimens Examined Total 69. Site 3 (21 year (Barbehenn et al., 1973). Cranial and msu-iit); Site 5 (4 msu-iit); Site 6 (1 msu-iit); Site 7 9 external measurements (Table 6) are comparable (4 fmnh); Site (2 dmnh); Site 10 (6 dmnh); to those of Dinagat, Leyte, and Mindanao (Mt. Site 11 (3 dmnh); Site 12 (6 dmnh); Site 13 (22 Kitanglad), but slight variations in most dimen- dmnh). sions are discernible (Heaney & Rabor, 1982; Order Carnivora Rickart et al., 1993; Heaney et al., unpubl. data). — Specimens Examined—Total 15. Site 3 (2 msu- Family Viverridae Civets iit); Site 7 (5 fmnh); Site 12 (1 dmnh); Site 13 (7 Paradoxurus hermaphroditus Jourdan, 1837 DMNH). The common palm civet occurs from Sri Lanka to Hainan and the Lesser Sunda Islands Rattus tanezwni Temminck, 1844 and is widespread in the Philippines (Heaney et Previously known in the Philippines as Rattus al., 1998). It has been recorded in agricultural rattus and Rattus the Oriental house mindanensis, and forested areas from sea level up to at least rat is a widespread nonnative rodent in the 2400 m (Heaney et al., 1991, 1999, unpubl. data). Philippines; it occurs from to Afghanistan We did not capture any palm civets, but in New Guinea, and Micronesia Indomalaya. March 1995 we were given a mandible from (Musser & Carleton, This 140 1993). 200-g rat a specimen taken by local hunters in the is most often found as a in urban and pest mountains above Mahinog, thus confirming the agricultural areas but can be common in earlier report by Gray (1843). A local hunter disturbed forest to 1800 m (Heideman et al., up identified P. hermaphroditus from pictures as 1987; et Heaney al., 1989, 1991, 1998, unpubl. being common on Camiguin.

42 FIELDIANA: ZOOLOGY — Specimens Examined Total 1.(1 fmnh). Fruit Bats—Because pteropodid bats lack sonar systems, they are easily captured in mist Viverra tangalunga Gray, 1832 nets. The 1960s surveys, which focused on birds, The Malay civet (or tangalung) is found from yielded many fruit bat specimens (Heaney, 1984), peninsular Malaysia to Sulawesi and is wide- most of them almost certainly from mist nets. spread in the Philippines (Heaney et al., 1998). It Those efforts obtained six species of fruit bats has been found in primary and secondary (Fig. 2). Our netting in 1992-1994 yielded no lowland, montane, and forest from sea mossy additional species but was not extensive. We level to at least 1700 m (Rickart et al., 1993; consider it quite possible that some additional Heaney et al., 1999). We did not obtain any on species may be present, especially Eonycteris Camiguin, but a local hunter identified Viverra spelaea and Rousettus amplexicaudatus, since tangalunga from photographs as being present both occur very widely in the Philippines on Camiguin; we tentatively accept this as a valid (Heaney et al., 1998), and possibly the large record. Acerodon jubatus and Pteropus vampyrus, which fly high above the canopy and so are difficult to Order Artiodactyla capture. However, our sampling was sufficient Suidae— Family Pigs that we consider it quite unlikely that the species Sus 1 886 philippensis Nehring, that are abundant and easily captured on Mind- The is a Philippine warty pig Philippine anao and associated islands (Heaney et al., 1989, endemic that occurs in the Greater Luzon, unpubl. data; Rickart et al., 1993) are present on Greater Mindanao, and Mindoro faunal regions; Camiguin, especially Haplonycteris fischeri and its numbers are declining (Oliver, 1992, 1999). It Ptenochirus minor and probably Megaerops formerly was abundant from sea level to at least ecaudatus. Suitable habitat for the high-elevation 2800 m in all habitats; now it is common only in specialist Alionycteris paucidentata is very limited remote forests (Heaney et al., 1991, 1999, in area on Camiguin, if present at all; although unpubl. data; Oliver, 1992, 1999; Garcia & we did limited netting in that habitat, we think it Deocampo, 1997). very unlikely that this species is present. On Camiguin in March 1995, we observed Sampling along the elevation gradient proba- hoof marks of wild pigs from disturbed lowland bly has produced a partial picture of variation in forest at 1 000 m up to primary montane forest at species richness (Fig. 2) but as noted may be 1275 m elevation (Site 7). Near the sampling site incomplete because our netting in 1992-1995 was at 1475 m (Site 8), we saw an active pig nest, and not extensive (Table 3). scattered pig trails were in clear evidence. A local Insectivorous Bats—Insectivorous bats use hunter said that they were commonly hunted and sophisticated sonar systems to navigate. They are often sold in a small local market at Owakan, thus difficult to capture in mist nets, which were Mahinog Municipality, but not in coastal cities. our only means of capturing them; further, our We were given two adult mandibles by hunters mist-netting efforts in 1992-1995 were limited in extent Table from pigs captured in forest in the early 1 990s in (see Heaney & Tabaranza, 2006a; 3). the mountains above Mahinog. For this reason, we believe that our sample of six Specimens Examined—Total 2. Above Mahi- species is likely to be quite incomplete and is not usable for estimates of as is nog (2 fmnh). species richness, often the case (Heaney et al., 2002). Our data contribute, however, to general knowledge of the natural history of these poorly known animals. Analysis and Discussion Nonvolant Small Mammals—Shrews, ro- dents, and other small mammals that can be Adequacy of Sampling: What Is Present and What captured in traps were poorly sampled by the

Is Absent? 1 960s field teams, which focused their efforts on birds (and secondarily on fruit bats). During Before interpreting field data, it is necessary to 1994 and 1995, we accumulated 2783 trap-nights evaluate the extent to which they are complete at six sites that included all major habitat types and reliable. In doing so here, we follow the on the island, yielding 188 captures (Table 1). A procedures used by Heaney et al. (1989, 1991, species-accumulation curve for these data (Fig. 3) 1999) and Rickart et al. (1993). shows that the number of species recorded for

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND No. Species

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

No. Trap-Nights

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

the concordance between our the island reached a peak of eight species obtained and and the information received from (including five nonnative species) by the end of specimens we conclude that deer are sampling at the second site (1055 trap-nights) local hunters. Thus, the smaller but and showed no further gain as we approached absent, along with conspicuous and the total of nearly 2800 trap-nights. Local species such as tarsiers, squirrels, flying hunters either were unfamiliar with squirrels or lemurs. knew them from Mindanao; given their conspic- uousness (Heaney et al., unpubl. data), they are Elevational Patterns very unlikely to be present. We conclude that the us from island is unlikely to support other species of Though our limited netting prevents small mammals that are present on Mindanao, commenting on the patterns of diversity and including gymnures (Podogymnura), tree shrews abundance of bats along the elevational gradient (Urogale), squirrels, or murid rodents (e.g., on Camiguin, our trapping data (Table 1) pro- Batomys, Limnomys, and Tarsomys). Further, vide a reasonably complete picture of the general the sampling along the elevational gradients was patterns among nonvolant species. The highest sufficiently intense that we are likely to have an density we encountered was at Site 3, in accurate assessment of the pattern of species agricultural land at ca. 150 m. However, these richness above about 800 m, although it must be were all nonnative rats and a shrew, representing noted that extensive habitat destruction at lower only three species, a pattern that we have often elevations may have had an extensive impact on seen (e.g., Heaney et al., 1989; Rickart et al., the current patterns (discussed below). 1991). The five native small mammal species Medium and Large Mammals—We collected were all present (or suspect to be present) from information only on the presence and absence of 1000 to 1300 m and declined in species richness large mammals and a mixed set of medium-sized to two species near the peak of Mt. Timpoong at species (none of which are readily captured in Site 8 (Table 1). Density (measured as the traps such as the ones we used), not on detailed number captured per 100 trap-nights) more than habitat use. We observed carefully during all our doubled from 1000 to 1275 m, then fell to a much time in the field for any evidence of additional lower level near the peak at 1475 m. However, species, and we interviewed many residents of the much of the change in density was driven by the islands and showed them photos of potential remarkably large numbers of Suncus murium at species; some of these residents are active the upper elevations; they were not taken at 1000 hunters. Neither we nor our informants had or 1200 m (though they were taken at 150 m), any evidence for tarsiers, flying lemurs, or deer were fairly common at 1300 and 1475 m, and on the island, though many people knew of deer were exceptionally abundant at 1275 m, where and tarsiers on Mindanao, and a few knew of we often saw and heard them during the day. lemurs. it flying squirrels and flying We consider These patterns for species richness, though unlikely that species other than those listed here limited in extent, are similar to those from are the voucher present given specimens we elsewhere in the Philippines, where species

44 FIELDIANA: ZOOLOGY richness is generally low at low elevations, ogale), flying lemurs (Cynocephalus), tarsiers increases to about the area of transition from (Tarsius), tree squirrels (Sundasciurus), pygmy montane to mossy forest, and declines with squirrels (Exilisciurus), flying squirrels (Peti- increasing elevation in mossy forest (Rickart et nomys), a murid (Tarsomys echinatus), and deer al., 1991; Heaney, 2001). The pattern of overall (Cervus). Among the murid rodents, Batomys is abundance is also similar to that documented also absent though often found in montane elsewhere, except that Suncus murinus is re- forest on Mindanao and associated islands markably abundant in the high-elevation mon- (Rickart et al., 1993). Limnomys bryophilus, L. tane and mossy forest, where usually it is absent sibuanus, and Tarsomys apoensis also occur on (Heaney, 2001). We have previously noted the Mindanao but only above about 1900 m eleva- abundance of S. murinus in similar habitat only tion (Musser & Heaney, 1992; Heaney et al., once, on Negros Island (Heaney et al., 1989), 1998; Rickart et al., 2003). which is notably depauperate of small mammals Those species of nonvolant mammals that are (Heaney, 1986). This raises a question: Is present on Camiguin form a group that is quite Camiguin depauperate, relative to its area, in consistent in one respect. Not all lowland species comparison to other islands in the Philippines? from Mindanao are present, but all those that This question is addressed in the next section. are present on Camiguin are among the relatively few species that are common in the lowlands of Biogeographic Implications Mindanao—or were present before human de- struction of lowland rain forests. As shown by Bats—Because our focus in conducting field Musser and Heaney (1992), Heaney (2001), and studies concentrated primarily on nonvolant Heaney et al. (unpubl. data), Crocidura, Apomys, mammals, we note only that we are confident Bullimus, Crunomys, Rattus everetti, Paradox- that the fruit bat community on Camiguin is at urus, Viverra, and Sus are among the few least somewhat depauperate relative to similar mammals that occur (or occurred until recently) elevations on Mindanao and associated islands. on Mindanao in lowland forest. With only one At least two species usually common at lower exception (Tarsomys echinatus), all those low- and middle elevations, Haplonycteris fischeri and land species of Mindanao noted above as being Ptenochirus minor, are almost certainly absent. absent from Camiguin are arboreal species. In Both of these are primarily associated with old- other words, the nonvolant mammal fauna of growth rain forest and generally do not travel Camiguin is composed only of nonarboreal small long distances out from under the canopy of mammals from Mindanao (or are their sister forest (Heaney et al., 1989, 1998; Heideman & taxa) and all the lowland larger mammals (all of that Heaney, 1989; Rickart et al., 1993). Some which are also not arboreal). We conclude additional species probably also are absent the mammalian fauna of Camiguin is highly (Alionycteris paucidentata and Megaerops wet- biased; that is, it is composed entirely of species morei); they also are associated with old-growth shared with and/or derived from Mindanao, but rain forest, though A. paucidentata in a distinctive it is not a random sample of the Mindanao high-elevation habitat (M. wetmorei is poorly fauna; rather, it is comprised of species that known). This level of reduction in species occur (or occurred before deforestation) on the in the richness is striking in view of the tight correlation lowland forest floor of Mindanao, not in the or between species richness and island area pre- forest canopy and not montane mossy viously found (Heaney, 1991a). However, the forest. uncertainty regarding the potential absence of Does the absence of arboreal mammals mean Eonycteris and Rousettus leads us to be cautious that Camiguin has a species-poor nonvolant in defining the extent to which species richness is mammal fauna relative to other islands in the low. Philippines? Perhaps surprisingly, the answer is Nonvolant Mammals—The data are more clearly no. With nine native nonvolant mam- falls almost on the certain regarding both small mammals (shrews mals, Camiguin precisely the islands that made and murid rodents) and the medium and larger same species-area curve as Mindanao mammals (deer, squirrels, and so on). It seems up Greater (Mindanao, Leyte, Bohol, While the fauna is biased certain that many genera present in the lowlands Biliran, and Maripipi). and in montane habitats on Mindanao and toward lowland, ground-living murid rodents, associated islands are absent: tree shrews (Ur- and small omnivorous carnivores, the species

HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND 45 was much richness is not reduced relative to islands that (lsnm). The manuscript improved by from P. D. Heideman and E. A. were recently connected to Mindanao itself comments Rickart. We thank Lisa Kanellos for preparation (Heaney. 1986; Rickart et al., 1993). However, was the the nonvolant small mammal community is of the figures. Funding provided by Marshall Field Fund. Ellen Thorne Smith Fund, apparently sufficiently species-poor to allow a nonnative shrew (Suncus minimis) to invade and the Barbara Brown Fund for Mammal Field Museum of Natural primary montane forest. The presence of this Research of the shrew in similar habitat has been noted on History. 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. Literature Cited 1993). We predict that S. minimis will be found in montane forest on other islands in the primary Balete, D. S.. B. R. Tabaranza, Jr., and L. R. Philippines with five or fewer native shrews and Heaney. 2006. An annotated checklist of the land rodents. birds of Camiguin Island, Philippines, pp. 58-72. In L. R. Heaney. ed.. The Mammals and Birds of Island. a Distinctive Center Conservation and Management Camiguin Philippines, of Biodiversity. Fieldiana Zoology, n.s., 106: 1-72. BARBEHENN, K.. J. P. SUMANGIL, AND J. B. LlBAY. 1973. As noted and Tabaranza (1997, by Heaney Rodents of the Philippine croplands. Philippine the native mammal fauna of is 2006a), Camiguin Agriculturalist, 56: 217-242. survival of dependent on the continued good- ESSELSTYN, J. A.. P. WlDMANN. AND L. R. HEANEY. quality forest at all elevations. The two mammal 2004. The mammals of Palawan Island. Philippines. Proceedings of the Biological of species unique to Camiguin, in particular, Society Washing- ton. 117: 271-302. apparently depend on forest with little or no Fooden. J. 1995. review of Southeast Asian disturbance, most of which currently occurs Systematic longtail macaques. Macaca fascicularis (Raffles. in steep areas above 800 m elevation. These [1821]). Fieldiana Zoology, n.s., 81: 1 206. forested areas are also crucial to the well-being Garcia. M. G. P., and A. H. Deocampo. 1997. Wild and of the human for stability population, pig meat trading in Iligan City, Mindanao. Sylva- they are the source of water for the island and trop. 5(1995):29-34. protect the lowlands from potentially devastat- Goodman. S. M., and N. R. Ingle. 1993. Sibuyan ing floods and landslides. Additionally, our Island in the Philippines-threatened and in need of conservation. 27: 174-180. observations indicate that the medium and large Oryx, Gray, J. E. 1843. List of the of mammals in mammals have been depleted by overhunting; specimens the collection of the British Museum. British these require protection if they are to survive. Museum (Natural History). The people of Camiguin benefit both personally Hall, R. 1998. The plate tectonics of Cenozoic SE and economically from the beautiful landscape Asia and the distribution of land and sea, pp. 99- and seashores of the both the island, through 132. In R. Hall and J. D. Holloway, eds.. Bio- tourism they make possible and from the geography and Geological Evolution of SE Asia. Publishers. Leiden. environmental stability they engender. Protect- Backhuys

— . 2002. Cenozoic ing the forests will benefit the people and wildlife geological and plate tectonic evolution of the SE Asia and the SW Pacific: equally. Computer-based reconstructions, model and anima- tions. Journal of Asian Earth Sciences, 20: 353^31. Heaney, L. R. 1984. Mammals from Camiguin Island, Philippines. Proceedings of the Biological Society of Acknowledgments Washington, 97: 119-f25.

— . 1986. Biogeography of mammals in Southeast Our thanks go to the Protected Areas and Asia: Estimates of rates of colonization, extinction, and Wildlife Bureau of the Philippine Department of speciation. Biological Journal of the Linnean Environment and Natural Resources, which Society. 28: 127-165. 1991a. An of of distribution provided encouragement and permits, and the analysis patterns and species richness among Philippine fruit bats Region 10 office of the DENR for logistical (Pteropodidae). Bulletin of the American Museum for were support. Specimens comparison kindly of Natural History. 206: 145-167. loaned by G. Hess (dmnh) and P. Myers (ummz) 1991b. A synopsis of climatic and vegetational and M. L. and H. Carleton, Gordon, Kafka change in Southeast Asia. Climatic Change, 19: 53-61.

46 FIELDIANA: ZOOLOGY . 1993. Biodiversity patterns and the conserva- of Camiguin Island, Philippines, a Distinctive Cen- tion of mammals in the Philippines. Asia Life ter of Biodiversity, Fieldiana Zoology, n.s., 106: Sciences, 2: 261-274. 1-72.

2000. Dynamic disequilibrium: A long-term, 2006b. A new species of forest mouse, genus large-scale perspective on the equilibrium model of Apomys (Rodentia, Muridae) from Camiguin island biogeography. Global Ecology and Biogeog- Island, Philippines, pp. 14-27. In L. R. Heaney, raphy, 9: 59-74. ed., The Mammals and Birds of Camiguin Island, a Distinctive Center of 2001. Small mammal diversity along eleva- Philippines, Biodiversity, Fieldiana 106: 1-72. tional gradients in the Philippines: An assessment of Zoology, n.s., patterns and hypotheses. Global Ecology and Bio- Heaney, L. R., E. K. Walker, B. R. Tabaranza, Jr., geography, 10: 15-39. and N. R. Ingle. 2002. Mammalian diversity in the 2004. Conservation biogeography in an Philippines: An assessment of the adequacy of current data. oceanic archipelago, pp. 345-368. In M. V. Lomo- Sylvatrop, 10(2000):6-27. lino and L. R. Heaney, eds., Frontiers of Bio- Heideman, P. D. 1989. Delayed development in Directions in the geography, New Geography of Fischer's pygmy fruit bat, Haplonycteris fischeri, in Nature. Sinauer Associates, Sunderland. the Philippines. Journal of Reproduction and Heaney, L. R., D. S. Balete, L. Dolar, A. C. Alcala, Fertility, 85: 363-382. P. C. N. R. M. A. Dans, Gonzales, Ingle, Lepiten, . 1995. Synchrony and seasonality of reproduc- L. R. P. S. E. A. B. R. W. Oliver, Ong, Rickart, tion in tropical bats. Symposium of the Zoological Tabaranza, Jr., and R. C. B. Utzurrum. 1998. A Society of London, 67: 151-165. of the mammalian fauna of the Philippine synopsis Heideman, P. D., J. A. Cummings, and L. R. Heaney. Islands. Fieldiana Zoology, n.s., 88: 1-61. 1993. Reproductive timing and early embryonic Heaney, L. R., D. S. Balete, E. A. Rickart, R. C. B. development in an Old World fruit bat, Olopteropus Utzurrum, and P. C. Gonzales. 1999. Mammalian cartilagonodus (Megachiroptera). Journal of Mam- diversity on Mt. 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48 FIELDIANA: ZOOLOGY A New Species of Hanging-Parrot (Aves: Psittacidae: Loriculus) from Camiguin Island, Philippines

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

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 curator and Philippine expert (e.g., Rand & Rabor 1960, 1969), had penciled the notation The Philippine Hanging-Parrot or Colasisi "subsp. nov" on the tag of a specimen from {Loriculus philippensis) has ten described sub- Camiguin in the fmnh collection. He never species distributed throughout the islands of the published a description. Here, we quantitatively evaluate the external and Philippines (Dickinson et al., 1991; Collar, 1997; morphology compare the color of referred to L. Juniper & Parr, 1998; Kennedy et al., 2000). The plumage specimens p. from with L. from subspecies L. p. apicalis has been reported to apicalis Camiguin p. apicalis of other of occur on the islands of Bazol, Balut, Camiguin, Mindanao and specimens subspecies L Our results demonstrate that the Dinagat, Mindanao, and Siargao (Fig. 1). How- philippensis. of L. is ever, Austin Rand, a former Field Museum Camiguin population p. apicalis separa- ble from all other populations of L. philippensis in It is further from all 1 plumage. separable Field Museum of Natural History, 1400 South in size. We Lake Shore Drive, Chicago, IL 60605-2496, U.S.A. neighboring populations body argue 2 that these differences warrant designating this Biological Sciences, University of Illinois at population as a distinct species and not a sub- Chicago, Chicago, IL 60607, U.S.A. of L. Similar have 3 species philippensis. arguments Current address: Division of Vertebrate Zoology- been made in separating L. bonapartei of the Ornithology, American Museum of Natural History, Sulu from the L. com- Central Park West at 79th Street, New York, NY Archipelago philippensis 10024-5192, U.S.A. plex (Juniper & Parr 1998). We present a formal

49 FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, PP. 49-57 Fig. 1. Map of the central and southern Philippines showing the locations of islands referred to in the text.

50 FIELDIANA: ZOOLOGY of the new taxon based description on plumage localities on Camiguin Island), 18 June 1968, and of adult males. morphology collected by D. S. Rabor and W. Sanguila. Diagnosis—A Loriculus hanging-parrot with characteristics of the philippensis group (see Front In contrast to other members of Methods Plate). this group, L. camiguinensis is characterized by a lack of sexual dimorphism in plumage color- All qualitative color comparisons were made ation. The overall color pattern of L. camigui- under natural light. Color names follow Smithe nensis is most like females of L. p. worcesteri (1975), and each color name (capitalized) is from Bohol, Leyte, and Samar and L. p. apicalis followed by its number in J.F.D. parentheses. from Mindanao but differs as follows: (1) the measured tarsus tail wing chord, length, length scarlet of the crown of L. camiguinensis does not of insertion of central rectrices to (from point tip extend as far onto the bright olive green nape as of longest rectrix), culmen bill it in length, length does both males and females of L. p. apicalis anterior of bill (from edge nostril), height (at and L. p. worcesteri; this pattern differs from that anterior of bill width anterior edge nostril), (at of chrysonotus, siquijorensis, regulus, bournsi, of and width with to edge nostril), gape calipers mindorensis, and philippensis, in which the scarlet the nearest 0.05 mm. were Specimens measured crown is highly reduced. (2) The width of the to avoid randomly any investigator bias. scarlet crown in L. camiguinensis narrows at the All statistical analyses were carried out using rear edge instead of being rounded as in all other the Statistica program (Statsoft Inc. 1995). populations. (3) The scarlet (sometimes orange) Mensural data were tested for normality using throat patch that is typical of males in L. Kolmogorov-Smirnov tests and Lilliefors prob- philippensis is lacking in L. camiguinensis; five abilities prior to all the analyses. Mensural of the Camiguin males had data on gonadal differences between males and females within development, reporting enlarged or slightly each study population were evaluated using one- enlarged testicles, which gives an indication of way analyses of variance (ANOVAs). We used their adult condition. (4) The face of L. both univariate (one-way ANOVAs and Bonfer- camiguinensis is extensively turquoise blue and roni post hoc tests) and multivariate analysis differs from that of females of L. philippensis (principal components and discriminant func- subspecies in that the blue of the face is darker tion) on log-transformed data to test for and more extensive, extending over the eye and mensural differences between specimens from onto the throat. (5) The turquoise blue in the Camiguin and those from neighboring popula- thighs of L. camiguinensis is darker than that tions of L. philippensis. Multivariate analyses of females of L. philippensis populations. (6) were used to reduce the dimensionality of data The blue in the inner edges of the rectrices above and facilitate the analysis of morphology in two and throughout below is darker in L. camigui- or three dimensions (Pimentel, 1979); we used the nensis. (7) Mean wing chord and tail length of varimax raw method to rotate the three compo- males and tail length of females of L. camigui- nensis are than of nents that are reported in the principal compo- significantly longer those L. 1 and nents analysis in order to improve interpretabil- nearby philippensis subspecies (Tables The overall is a darker ity of the resulting patterns. Collecting localities 2). (8) green plumage shade with less of a on are described by Heaney and Tabaranza (2006a). yellowish tinge, especially the back. —General Loriculus camiguinensis, new species Description of Holotype plumage Parrot Green (160) with slightly orange tinge in Camiguin Hanging-Parrot the upperparts, more yellowish tinge on under- pays; forehead and forepart of crown Scarlet Holotype—Field Museum of Natural History (14) fading to orange at rear-edge; thin Orange No. 284389, adult male from Kasangsangan, Yellow (18) band on nape; lores, chin, cheek, and Municipality of Catarman, Camiguin Province, throat closest to Turquoise Blue (65); rump and Camiguin Island, Philippines, elevation between upper tail-coverts scarlet; Turquoise Green (64) 1000 and 2000 ft (300-600 m; approx. 9°11'N, markings on the sides of the rump; thighs slightly 124°40'E; see Heaney & Tabaranza, 2006a, for paler Turquoise Blue; rectrices Emerald Dark more explanation of this and other collecting 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; Mati: Mt. Mayo, Limot (male: fmnh 277864); rectrices dark Cerulean Blue below; flight feath- Misamis Oriental Province: Manticao: Tuod, ers black above with Emerald Dark Green outer Camp Dundue (male: fmnh 283788; female Province: edges, flight feathers black below with inner fmnh 283787); Misamis Oriental Opol: edges with extensive Cerulean Blue; greater Malubato (male: fmnh 283785); South Cotabato: underwing-coverts Cerulean Blue and lesser un- Tupi: Mt. Matutum (male: fmnh 279330); derwing-coverts closest to Spectrum Green (62). Bukidnon Province: Malaybalay, Mt. Katanglad Soft part colors of dried specimen: upper (male: fmnh 262475, 262476; female: fmnh mandible closest to Spectrum Orange (17) at 262474); Bukidnon Province: Lantapan: Ko- base grading to yellow with Gray Horn (91) at toon, Mt. Katanglad SE slope (male: dmnh tip and along tomia; lower mandible with similar 2983); Lanao Norte Province: Iligan City, pattern, but orange at the base; cere Grayish- Mainit, Mahayahay (female: fmnh 283786); Horn; feet and tarsi, yellow-horn. Surigao del Sur Province: Car-Can-Mad-Lan Measurements of Holotype (mm) —Wing area (female: fmnh 275002); Zamboanga del Sur (99.8), tarsus (11.7), tail (49.5), culmen (18.0), Province: Zamboanga (male: dmnh 36993); bill length (15.0), bill height (11.1), bill width Davao Oriental Province: Sigaboy (males: dmnh (6.4), gape width (8.2). 36224, 36226); Davao del Sur Province: Padada Distribution—Loriculus camiguinensis is (male: dmnh 36233). known only from the forests of Camiguin Island. L. p. worcesteri (11 males, 8 females): Bohol Specimens have been collected between 1000 and Island: Bohol Province: Sierra Bullones (males: 4500 ft (300-1350 m) in the municipalities of fmnh 223025, 223026, 223029, 223030, 223034, Catarman and Mahinog (Balete et al., 2006; 223036, 223037; females: fmnh 223027, 223327, & Heaney Tabaranza,— 2006a). 223028, 223033, 223035, 223039); Leyte Island: Etymology We name this species after the Leyte Province: Burauen, Buri, Ma-Alngon Philippine Island of Camiguin, to which this (male: fmnh 276302; female: fmnh 276300); species appears to be endemic. Leyte Province: Burauen, Buri, Mt. Lobi range, Specimens Examined—We examined the fol- Tambis (male: fmnh 276299; female: fmnh lowing specimens from Field Museum of Natural 276298); Samar Province: Mt. Capotoan (male: History (fmnh) and Delaware Museum of fmnh 247411); Western Samar Province: Matu- Natural History (dmnh): guinao (male: fmnh 247410). L. camiguinensis (Camiguin Island) (19 males, L. p. regulus (Negros Island) (5 males, 4 4 females, all known specimens of this new females): Negros Oriental Province: Bayawan, taxon): Camiguin Province: Catarman Munici- Basay (male: fmnh 257121); Negros Oriental pality; Kasangsangan (males: fmnh holotype, Province: Santa Catalina, Inubungan (male: fmnh 284391, 284392, 284393; dmnh 19950, fmnh 219314; male: fmnh 188579); Negros 19958, 19960, 19961, 19962: females fmnh Oriental Province: Sicopon River (male: fmnh 284390; dmnh 19959); Camiguin Province: Cat- 185483); Negros Oriental Province: Amio (males: arman Municipality; Catarman Mountain (male: fmnh 188545, 188548; females: fmnh 188544, dmnh 19949); Camiguin Province: Mahinog 188553); Negros Oriental Province: Pamo-at Municipality; Matugnao, Mt. Timpoong (males: (male: fmnh 188550). fmnh 286742, L. 286743, 286744, 286745; dmnh p. chrysonotus (captive specimen, presum- 19951, 19952, 19953, 19954, 19965; females: ably from Cebu Island) (1 male): Cebu Province: fmnh 286746, 286747). Exact locality unknown (fmnh 252666). L. p. (Mindanao 5 L. apicalis Island) (19 males, p. siquijorensis (Siquijor Island) (1 male): North Cotabato females): Province: Mt. Apo, Siquijor Province: San Juan: Tag-ibo (fmnh Todaya (male: fmnh 184090); North Cotabato 222741). Province: Mt. Apo, Galog (male: dmnh L. bournsi 36227); p. (Sibuyan Island) (1 male, 1 Agusan del Norte: Mt. Lewed Hilong-Hilong, female): Romblon Province: Goangan, 3 km SE (male: fmnh Misamis 275003); Occidental: Zam- Magdiwang (male: fmnh 358288); Romblon boanga Peninsula, Mt. Gandwan Province: Malindang, Exact locality unknown (female: fmnh (males: fmnh 227136, 227138, 227139; female: 11081). fmnh 227137); Misamis Occidental: L. Zamboanga p. mindorensis (Mindoro Island) (1 male, Peninsula. Mt. Malindang, Masawan (males: 1 female): Oriental Mindoro Province: fmnh 227134, 227135); Davao Oriental Province; Calapan (male: fmnh 19927); Occidental Mind-

52 FIELDIANA: ZOOLOGY la rsi no o 03 on •? K*S ^Ci.

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3 3 Table 2. Univariate statistical comparisons be- ol' L. tween /.. camiguinensis and adjacent populations = = 1 L. 2 L. philippensis. Populations ( camiguinensis: = 4 = p. apicalis; 3 L. p. worcesteri; and 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 o

L camiguinensis O L p. apicalis o L p. worcesteh A 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.

most commonly involves undeveloped gonads, neighboring populations. We were able to make and there are clearly labeled tags indicating direct comparisons with specimens of all sub- that the gonads of some specimens were de- species except L. p. dohertyi (Basilan). It is veloped. If mistakes were made, the plumage possible that L. camiguinensis is more closely characters of the specimens described above related to some parts of the L. philippensis group would logically argue that all males were than others, which would make L. philippensis misidentified as females. Again, this seems highly paraphyletic (Funk & Omland 2003). Despite unlikely given the number of males (19). this possible relationship to the widespread L. Furthermore, it seems highly unlikely that philippensis group, we believe L. camiguinensis a series of 23 randomly collected parrots would sufficiently differentiated to be beyond concerns all have been females. Certainly, there is nothing expressed about recognizing new species based like this in other series of Loriculus collected by on minor morphologic differences (e.g., Collar et Rabor in other parts of the Philippines. Thus, al., 1999). Based on geographic distance and the while we cannot say that all specimens are overall pattern of plumage coloration, L. cami- unequivocally identified to sex correctly, we feel guinensis most closely resembles populations of that adults of both males and females are L. p. apicalis and L. p. worcesteri, but no included in these series. 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 Discussion populations to lose bright plumage, leading to a lack of sexual dichromatism (see references in Our results demonstrate that L. camiguinensis Peterson, 1996); L. camiguinensis is the only is diagnosable from populations of L. philippen- member of the genus without sexual dichroma- sis in plumage. It also differs in size from all tism in plumage.

TELLO ET AL.: A NEW SPECIES OF HANGING-PARROT The recognition of this distinctive taxon et al.. 2001) and another form, L. p. siquijorensis, coincides with recent surveys of the small may be extinct as well (Forshaw, 1989; Kennedy mammal fauna of Camiguin Island that have et al., 2000). The combined populations of discovered two new species of rodents (Rickart et Mindoro, Sibuyan, Negros, Panay, Tablas, al., 2002; Heaney et al., 2006). Camiguin is Romblon, Masbate, Ticao, Guimaras, and Basi- believed to be the smallest Philippine island to lan (including mindorensis, bournsi, regulus, and harbor endemic species of birds and mammals dohertyi) may total no more than 5000 individ- (Heaney & Tabaranza, 1997, 2006a). The island uals (Juniper & Parr, 1998). The current popu- has been continuously isolated from its large lation size of L. camiguinensis is not known (but southern neighbor, Mindanao, even during see Heaney and Tabaranza, 2006a, for an periods of low sea level during the "ice ages" assessment of remaining habitat on the island). of the Pleistocene, when sea levels dropped to Without field data on its status, we defer from 120 m below present levels (Heaney, 1986, suggesting how this species should be character- 1991a, 1991b; Fairbanks, 1989; Heaney and ized under international threat criteria (IUCN Tabaranza, 1997, 2005b; Heaney and Regalado, Species Survival Commission, 1994). However, 1998; Hanebuth et al., 2000), and this may have because Camiguin is a small island that has played a role in the differentiation of Camiguin's experienced extensive deforestation, the conser- fauna from that of Mindanao (Steppan et al., vation status of this newly described species 2003). clearly requires assessment. Field study is needed The value of museum collections is well to establish the population size and requirements illustrated with this description. These collec- as a prerequisite for conservation planning and tions were essential in the recognition and action. 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 Acknowledgments immature plumage or an error in sexing of We are to G. Hess a specimen (a female incorrectly identified as grateful of the Delaware Museum of Natural a male; see above). However, the presence of History for loan of speci- mens under his a series of specimens from different localities care and to M. Skakuj for (with data on gonadal development) has allowed painting the plate. Bruce Patterson and P. us to compile meaningful data sets on morpho- Wagner provided help with data analyses, and S. Bober in logical variation and assess within-population helped preparing Figure 1. The variation in color. manuscript was greatly improved by comments from D. S. This new species also illustrates the need for Balete, A. Carnaval, L. R. Heaney, S. P. C. additional taxonomic and systematic research on Ware, Rasmussen, and N. J. Collar, the Loriculus hanging-parrots to understand the although this does not imply that all agree with the status we evolutionary patterns in the group and to convey on this taxon. evaluate the possibility that some of the other allopatric forms of L philippensis may also deserve species status. The issue of assessing the Literature Cited taxonomic status of allopatric populations in the Philippines has long been as a chal- recognized Balete, D. S., B. R. Tabaranza, Jr., and L. R. for lenge conservationists (Collar et al., 1999; Heaney. 2006. An annotated checklist of the land birds of Peterson et al., 2000). To date, little attention has Camiguin Island, Philippines, pp. 58-72. In L. The been given to the conservation of Loriculus Heaney, R., ed., Mammals and plight Birds of For Camiguin Island, Philippines, a Distinctive parrots. instance, none was included by Center of Biodiversity. Fieldiana Zoology, n s 106: Collar et al. (1999) in their list of threatened 58-72. bird Philippine species. This lack of attention can Collar, N. J. 1997. Family Psittacidae (Parrots), be correlated with 280-477. In directly the designation of L. pp. del Hoyo, J., A. Elliott and J. as a Sargatal, eds., Handbook of the Birds of philippensis polytypic species. The plight of the World, Vol. 4. Lynx Edicions. Barcelona. these populations is cause for concern, as Collar. N. J., N. A. D. and B. R Loriculus p. chrysonotus from the island of Cebu Mallari, Tabaranza. Jr. 1999. Threatened birds of the is believed to be extinct (Forshaw, 1989; Mallari Philippines. Bookmark, Makati City.

56 FIELDIANA: ZOOLOGY E. R. S. Dickinson, C, Kennedy, and K. C. Parkes. pp. 28^8. In Heaney, L. R., ed.. The Mammals and 1991. The Birds of the An Philippines: Annotated Birds of Camiguin Island, Philippines, a Distinctive Check-List. British Union Check- Ornithologists' Center of Biodiversity, Fieldiana Zoology, n.s., 106: List No. 12. Tring, UK. 1-72. R. 1989. Fairbanks, G. A 17,000 glacio-eustatic sea IUCN Species Survival Commission. 1994. IUCN Red level record: Influence of rates on the glacial melting list categories, as approved by the 40th meeting of event Younger Dryas and deep-ocean circulation. the IUCN Council, Gland, Switzerland, 30 Novem- Nature, 342: 637-642. ber 1994. IUCN-The World Conservation Union, Forshaw, J. M. 1989. Parrots of the World. Lans- Gland, Switzerland. downe Melbourne. Editions, Juniper, T., and M. Parr. 1998. Parrots: A Guide to Funk, D. J., and K. E. Omland. 2003. Species-level Parrots of the World. Yale University Press, New paraphyly and polyphyly: Frequency, causes and Haven, Connecticut. with insights from animal mitochon- consequences Kennedy, R. S., P. C. Gonzales, E. C. Dickinson, H. drial DNA. Annual Review of and Ecology System- C. Miranda, Jr., and T. H. Fisher. 2000. A Guide atics, 34: 397^423. to the Birds of the Philippines. Oxford University Hanebuth, T., K. Stattegger, and P. M. Grootes. Press, Oxford. 2000. Rapid flooding of the Sunda Shelf: A late- Mallari, N. A. D., B. R. Tabaranza, Jr., and M. J. sea-level record. Nature, 288: 1033-1035. glacial Crosby. 2001. Key Conservation Sites in the L. R. 1986. of in Heaney, Biogeography mammals Philippines. Bookmark, Makati City. Southeast Asia: Estimates of rates of colonization, Peterson, A. T. 1996. Geographic variation in sexual extinction, and Journal of the speciation. Biological dichromatism in birds. Bulletin of the British Linnean Society, 28: 127-165. Ornithologists' Club, 116: 156-172. — . 1991a. An analysis of patterns of distribution Peterson, A. T., L. S. Ball, and K. W. Brady. 2000. and species richness among Philippine fruit bats Distribution of the birds of the Philippines: Bio- (Pteropodidae). Bulletin of the American Museum geography and conservation priorities. Bird Conser- of Natural History, 206: 145-167. vation International, 10: 149-167. -. 1991b. A synopsis of climatic and vegetational Pimentel, R. A. 1979. Morphometries: The Multivar- change in Southeast Asia. Climatic Change, 19: 53-61 . iate Analysis of Biological Data. Kendall/Hunt, L. and J. C. Jr. 1998. Heaney, R., Regalado, Dubuque, Iowa. Vanishing Treasures of the Philippine Rain Forest. Rand, A. L, and D. S. Rabor. 1960. Birds of the The Field Museum, Chicago. Philippine islands: Siquijor, Mount Malindang, Heaney, L. R., and B. R. Jr. 1997. A — Tabaranza, Bohol, and Samar. Fieldiana Zoology, 35: 221 preliminary report on mammalian diversity and 441. conservation status of Camiguin Island, Philippines. . 1969. New birds from Camiguin South, Sylvatrop (1995), 5: 57-64. Philippines. Fieldiana Zoology, 51: 157-168. . 2006a. Introduction to the mammal and land Rickart, E. A., L. R. Heaney, and B. R. Tabaranza, bird faunas of Camiguin Island, Philippines, pp. 1- Jr. 2002. Review of Bullimus (Muridae: Murinae) 13. In Heaney, L. R., ed., The Mammals and Birds and description of a new species from Camiguin of Camiguin Island, Philippines, a Distinctive Cen- Island, Philippines. Journal of Mammalogy, 83: ter of Biodiversity, Fieldiana Zoology, n.s., 106: 421^136. 1-72. Smithe, F. B. 1975. Naturalist's Color Guide. The -. 2006b. A new species of forest mouse, genus American Museum of Natural New York. Apomys (Rodentia, Muridae) from Camiguin History, Island, Philippines, pp. 14-27. In Heaney, L. R., Statsoft Inc. 1995. Statistica for Windows, Statsoft ed., The Mammals and Birds of Camiguin Island, Inc., Tulsa. a Distinctive of Philippines, Center Biodiversity. Steppan, S., C. Zawadski, and L. R. Heaney. 2003. A Fieldiana 106: 1-72. Zoology, new series, molecular assessment of phylogenetic relationships Heaney, L. R., B. R. Tabaranza, Jr., D. S. Balete, and patterns of phylogenesis in the Philippine murid and N. Rigertas. 2006. Synopsis and biogeography rodent Apomys. Biological Journal of the Linnean of the mammals of Camiguin Island, Philippines, Society, 80:699-715.

TELLO ET AL.: A NEW SPECIES OF HANGING-PARROT 57 An Annotated Checklist of the Birds of Camiguin Island, Philippines 3 1 3 2 Danilo S. Balete, ' Bias R. Tabaranza, Jr., and Lawrence R. Heaney

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 et al., 1999; Peterson et al., 2000; but see Mallari et al., 2001, Ong et al., 2002). We note that Although the avifaunas of many of the Camiguin Norte, which lies north of Luzon, is Philippine islands have been reported in com- often confused with Camiguin. As documented prehensive fashion (summarized by Dickinson et in the following list, the land birds of Camiguin al., 1991), no such report has been made for Island include a diverse assemblage of at least 57 Camiguin Island, with the result that Camiguin species. These records are based on voucher has not been included in most analyses of avian specimens that were obtained principally during and biogeography conservation priorities (e.g., the 1960s and 1990s and are deposited at the Hauge et al., 1986; Dickinson et al., 1991; Collar Delaware Museum of Natural History (dmnh), the Field Museum of Natural History (fmnh), 1 and State Laksambuhay Conservation Inc., 10241 Mt. Bu- Mindanao University-Iligan Institute lusan Street, U-2. Los Banos, Laguna, Philippines. of Technology (msu-iit), as detailed below and in 2 1 & Addi- Department of Biology, Uigan Institute of Tech- Chapter (Heaney Tabaranza, 2006a). nology, Mindanao State University, Iligan City, tional records based on specimens at the Lanao del Norte, Philippines. Museum of Comparative Zoology (mcz), Cam- 3 Field Museum of Natural History, 1400 South bridge, as cited in Dickinson et al. (1991), are Lake Shore Drive, Chicago. IL 60605-2496, U.S.A. included as well.

58 FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, PP. 58-72 The of 55 of land — presence species breeding Specimens Examined Total 1. Site 1 (1 birds on an island the size of Camiguin in the fmnh). Philippines is not remarkable; indeed, as docu- mented of birds below, surveys on Camiguin Family Accipitridae—Hawks and Eagles were not and the current total is exhaustive, well Spilornis cheela—Crested Serpent-Eagle below the total number to be likely present. The Crested Serpent-Eagle occurs from India that is an oceanic However, given Camiguin to China, Taiwan, Ryukyus, and Southeast Asia; island surrounded water but about by deep only it is found throughout the Philippines, except the 10 km from the shore of these data Mindanao, Babuyan and Batanes groups of islands, in forest are a useful indicator of the of these 55 ability and forest edge (Dickinson et al., 1991; Kennedy to cross a sea channel of this species readily et al., 2000). Two male specimens were taken on an extensive of time. breadth, given period 13 June 1968 on Mt. Catarman, one at 2000 ft Further, as noted the avifauna of below, (ca. 600 m; Site 9), the other at an unknown is now considered to include four Camiguin elevation. We often saw them at nearly all endemic of birds everetti catar- subspecies (Ixos elevations during fieldwork in 1994 and 1995, manensis, azurea Hypothymis catarmanensis, especially near forest. Diceum trigonostigma isidroi, and Zosterops Specimens Examined—Total 2. Site 9 (1 dmnh; and an endemic nigrorum catarmanensis) species 1 fmnh). of parrot {Loriculus camiguinensis; Tello et al., This observation indicates that 2006). genetic Family Megapodidae—Megapodes and Scrubfowl differentiation has taken in at least some of place [Megapodius cumingii—Tabon] these in one case to species, leading significant The Tabon ranges from Sulawesi and Borneo change. It is our hope that this paper will to the Philippines; once widespread throughout encourage more extensive and focused analysis the country, it is now found mainly in coastal of the avifauna of on both Camiguin, ecology scrub but also occurs in lowland to montane and evolution. forest (Dickinson et al., 1991; Kennedy et al., For notes on methods of collecting and for 2000). In an earlier enumeration of Philippine a list and description of the collecting sites, see birds, Camiguin was not identified as part of the Heaney and Tabaranza (2006a). As noted in that Tabon's range (Dickinson et al., 1991). However, elevations from the 1960s should be chapter, in the more recent compilation, Kennedy et al. viewed as rough estimates. Species whose names (2000) listed Camiguin as part of the range of are listed in brackets are of uncertain documen- this species without further reference to the tation and are not included in our of tally species source of their data. Thus, we have not included numbers. Unverified sight records are listed at M. cumingii in our tally of birds from Camiguin the end of this paper. We define "first records" but note its potential presence. as those not reported by Dickinson et al. (1991) and/or Kennedy et al. (2000). Family Phasianidae—Partridges, Pheasants, and Quail Gallus gallus—Red Junglefowl Red occurs from India to Accounts of Species The Junglefowl southern China and Southeast Asia, including the Family Ardeidae—Bitterns, Egrets, and Herons virtually all of the Philippines, except Ixobrychus cinnamomeus—Cinnamon Bittern Batanes group of islands, in forest and forest The Cinnamon Bittern ranges from India to edge up to 2000 m (Dickinson et al., 1991; China, Ryukyus, Taiwan, and Southeast Asia; in Kennedy et al., 2000). A male specimen was at 3150 ft 950 the Philippines it occurs on most islands, except taken on Mt. Timpoong (ca. m) male the Babuyan and Batanes group, in marshes, rice on 14 June 1969 (Site 13). In May 1994, a was taken in a Victor in fields, and wetlands (Dickinson et al., 1991; specimen trap Kital-is, at 1200-1400 m We also heard Kennedy et al., 2000). A single specimen was Sagay (Site 6). in 1995 close to our taken in a highly disturbed agricultural area in them March crowing Balbagon, Mambajao, on 28 May 1992, at 10 m, campsite on Mt. Timpoong at 1275 m (Site 7). — 2. 6 msu- the first record of this species from the island Specimens Examined Total Site (1 13 (Site 1). iit); Site (1 dmnh).

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 59 to about 1600 m Family Rallidae—Coots, Crakes, Rails, secondary forest up (Dickinson and Waterhens et al., 1991; Kennedy et al., 2000). It was recorded from Mt. Catarman and the Porzana fusca- Ruddy-breasted Crake nearby of at 1 000-4500 ft 300- The Ruddy-breasted Crake ranges from India area Kasangsangan, (ca. 1400 in June 1968 9 and Further to Japan, Ryukyus, and Southeast Asia; in the m) (Sites 11). records were obtained from Mt. at Philippines it has been recorded only on the Timpoong 3150-4000 ft (ca. 950-1200 m) and the nearby islands of Bohol, Cagayancillo, Leyte, Luzon, area of Puntod at 800 ft 250 in June 1969 Mindanao, Mindoro, Negros, Panay, Samar, (ca. m) 13 and 14, We also recorded and Sibuyan, usually in marshes and rice fields (Sites respectively). it in 1994 in Kital-is, at 900-1100 m but also along forest paths up to 1500 m May Sagay, (Site 4). (Dickinson et al., 1991; Kennedy et al., 2000). Specimens Examined—Total 12. Site 4 msu- A male specimen was taken in Kasangsangan in (1 iit); Site 9 (3 dmnh, 1 fmnh); Site 1 1 (3 fmnh); the vicinity of Mt. Catarman at 1000 ft (ca. Site 13 (2 Site 14 (2 fmnh). 300 m) on 22 June 1968 (Site 11). This is the first fmnh); record of this species on Camiguin. — — Ptilinopus occipitalis Yellow-breasted Fruit- Specimens Examined Total 1. Site 11 (1 Dove FMNH). The Yellow-breasted Fruit-Dove is endemic to

the it is Rallina eurizonoides—Slaty-legged Crake Philippines; fairly widespread through- out the except the Batanes, The Slaty-legged Crake occurs from India and country, Babuyan, Palawan, and Sulu of islands, in forest Sri Lanka to Taiwan, Ryukyus, and Southeast groups up to 1800 m (Dickinson et al, 1991; Kennedy et Asia. In the Philippines it is rather uncommon on It was recorded on Mt. Catarman and most islands but absent on the Palawan and Sulu al., 2000). the areas of and groups of islands, in scrub and primary to nearby Gidag-on Kasangsangan at 1000^1500 ft (ca. 300-1400 m) in June 1968 secondary forest up to 900 m (Dickinson et al., (Sites 9-11). Additional records came from Mt. 1991; Kennedy et al., 2000). The record on at 3150^4800 ft (ca. 950-1450 in Camiguin, a first for this island, consists of two Timpoong m) June 1969 (Sites 12 and 13). specimens, both taken in Victor traps, from Specimens Examined—Total 18. Site 9 Kital-is, Sagay, at 900-1100 m on 19 May 1994 (3 dmnh, 3 fmnh); Site 10 (1 dmnh, 1 fmnh); Site (Site 4) and at 1200-1400 m on 26 May 1994 1 1 (1 fmnh); Site 12 (1 fmnh); Site 13 (4 dmnh, 4 (Site 6). The following year, another was taken fmnh). on Mt. Timpoong in primary montane forest at 1275 m on 23 March 1995 (Site 7). Columba vitiensis—Metallic Pigeon Specimens Examined—Total 3. Site 4 msu- (1 The Metallic Pigeon occurs on Borneo, iit); Site 6 (1 msu-iit); Site 7 (1 fmnh). Sulawesi, Moluccas, Lesser Sunda Islands, New Guinea, and Samoa; it is an uncommon resident Family Columbidae—Doves and Pigeons of most islands throughout the Philippines, in Treron vernans—Pink-necked Green-Pigeon lowland to mossy forest (Dickinson et al., 1991; The Pink-necked is Green-Pigeon widespread Kennedy et al., 2000). Records from Camiguin in Southeast Asia; it has been recorded through were all taken in 1968. A female specimen was most of the Philippines, the and except Babuyan taken from Mt. Catarman at 2000 ft (ca. 600 m) Batanes of in lowland groups islands, principally on 14 June (Site 9) and three adult males and an forest below 300 m second-growth (Dickinson, adult female from Kasangsangan at 1000- 1991; et al., 2000). An adult male 2000 ft Kennedy (ca. 300-600 m) in the vicinity of Mt. was taken on 29 June 1969 in specimen Puntod, Catarman on 11-18 June (Site 11). Mahinog, at 800 ft (ca. 250 m). — Specimens Examined Total 5. Site 9 (1 Specimens Examined—Total 1. Site 14 (1 fmnh); Site 11 (3 dmnh, 1 fmnh). dmnii). Macropygia phasianella—Reddish Cuckoo-Dove Phapitreron leucotis—White-eared Brown-Dove The Reddish Cuckoo-Dove ranges from Bor- The White-eared Brown-Dove is endemic to neo, the Moluccas, Sulawesi, and the Lesser the Philippines; it occurs the throughout country Sunda islands to New Guinea and Samoa; it with the of the and exception Babuyan, Batanes, occurs throughout the Philippines, in lowland to Palawan groups of islands, in and forest primary mossy above 2000 m (Dickinson et al.,

60 FIELDIANA: ZOOLOGY et 1991; Kennedy al., 2000). It was recorded on Specimens Examined—Total 5. Site 11 (2 the Mt. Catarman and nearby area of Kasang- dmnh, 2 fmnh); Site 14 (1 fmnh). sangan at 1000-2500 ft (ca. 300-750 m) in June 1968 9 and Additional records (Sites 11). came Family Psittacidae—Lorikeets, Cockatoos, Parrots, from Mt. Timpoong at 3150-5700 ft (ca. 950- and Racquet-tails 1700 and the area of Puntod at 800 ft m) nearby Loriculus camiguinensis—Camiguin Hanging- (ca. 250) in June 1969 (Sites 12-14). In May Parrot 1994, we recorded it in Kital-is, at 1000- Sagay, Described as a new species in this volume 1400 m 5 and 6). (Sites (Tello et al., 2006), this is the only species of bird Specimens Examined—Total 30. Site 5 msu- (1 currently recognized as endemic to Camiguin. Its Site 6 Site 9 Site 1 1 iit); (2 msu-iit); (1 fmnh); (3 elevational range, based on specimens collected dmnh, 3 Site 12 dmnh, 7 Site 13 fmnh); (9 fmnh); in the 1960s, is 300 to 1350, thus including 2 Site 14 (1 dmnh, fmnh); (1 fmnh). lowland and montane rain forest. Most of this habitat has been destroyed on Camiguin, leading Streptopelia bitorquata—Island Collared-Dove to concern for its conservation status. The Island Collared-Dove is found from Specimens Examined—Total 23. Site 9 (2 Borneo and the Lesser Sunda islands to Java; dmnh); Site 11 (5 dmnh, 5 fmnh); Site 13 (5 in the Philippines it is a fairly uncommon dmnh, 6 fmnh). resident of most islands, except the Batanes group of islands, mainly in relatively open fields Family Cuculidae—Cuckoos, Malkohas, and Coucals in the lowlands and sometimes in mangrove Cacomantis variolosus—Brush Cuckoo (Dickinson et al., 1991; Kennedy et al., 2000). The Brush Cuckoo occurs from Southeast Two male specimens were collected from Pun- Asia to Australia and the Southwest Pacific; it is tod, Mahinog, at 700-800 ft (ca. 200-250 m), found throughout the Philippines, except the one of which was a juvenile taken at 800 ft (ca. and Batanes of islands, from 250 m) on 28 June 1968 (Site 14). Babuyan groups mangrove to mossy forest at 2000 m (Dickinson Specimens Examined—Total 2. Site 14 (1 et al., 1991; et al., 2000). The records dmnh, 1 fmnh). Kennedy — from Camiguin were taken from Mt. Catarman Geopelia striata Zebra Dove at 2500 ft (ca. 750 m) in June 1968 (Site 9) and The Zebra Dove ranges from Southeast Asia from Mt. Timpoong at 3150^1800 ft (ca. 950- to Australia; it occurs throughout the Philip- 1450 m) in June 1969 (Site 13). We also recorded pines, except the Babuyan and Batanes groups of it in Kital-is, Sagay, at 900-1100 (Site 4) and islands, in open country and cultivated areas in 1200-1400 m in May 1994 (Site 6). the lowlands (Dickinson et al., 1991; Kennedy et Specimens Examined—Total 7. Site 6 (1 msu- us netted one al., 2000). One of (B.R.T.) some iit); Site 9 (1 fmnh); Site 13 (3 dmnh, 2 fmnh). 50 m away from the beach in Manuyog, Sagay, scolopacea—Common Koel on 10 May 1994 (Site 3), but it was released. No Eudynamys The Common Koel occurs from India other specimen was recorded of this species in through Southeast Asia to northern it is found 1994. It was not recorded during our fieldwork in Australia; 1992 and 1995. throughout the Philippines, in primary and Specimens Examined—None. secondary lowland forest (Dickinson et al., et The — 1991; Kennedy al., 2000). single specimen Chalcophaps indica Common Emerald-Dove from Camiguin was taken on Mt. Timpoong at is found The Common Emerald-Dove from 3150 ft (ca. 950 m) on 17 June 1969 (Site 13). India and Sri Lanka to China, Taiwan, Ryukyus Specimens Examined—Total 1. Site 13 (1 and Southeast Asia, New Guinea, and Australia; fmnh). it occurs throughout the Philippines, often in — Coucal second-growth forest in the lowlands up to Centropus viridis Philippine The Coucal is endemic to the 1000 m (Dickinson et al., 1991; Kennedy et al., Philippine where it occurs the 2000). Records on Camiguin consist of speci- Philippines, throughout mens taken in Kasangsangan, near Mt. Catar- country, except the Palawan group of islands, to forest at 2000 m et man, at 1000-2500 ft (ca. 300-750 m) in June from grassland (Dickinson et The records on 1968 (Site 11) and from Puntod at 800 ft (ca. al., 1991; Kennedy al., 2000). consist of taken from 250 m) in May 1969 (Site 14). Camiguin specimens

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 61 in June at at 1000-2000 ft (ca. 300-600 m) Kasangsangan in the vicinity of Mt. Catarman gan at 1968 1968 (Sites 9 and 11) and on Mt. Timpoong 1000 1500 it (ca. 300 450 m) in June (Site 3150-4800 ft (ca. 950-1450 m) in June 1969 11) and from Mt. Timpoong at 3150 ft (ca. 12 and 950 m) in June 1969 (Site 13). (Sites 13). Examined—Total 31. Site 9 (1 Specimens Examined—Total 6. Site 11 (1 Specimens Site 11 dmnh, 7 fmnh); Site 12 (2 dmnii. 2 fmnh); Site 13(1 dmnh, 2 fmnh). fmnh); (2 13 9 dmnh, 1 fmnh); Site (9 dmnh, fmnh).

Family Strigidae—Owls Swiftlet Ninox philippensis Philippine Hawk-Owl Collocalia troglodytes—Pygmy the is endemic to the The Philippine Hawk-Owl is endemic to The Pygmy Swiftlet Philip- of the it occurs most of the Philippines, occurring throughout most pines, where throughout and Sulu islands, except the Babuyan. Batanes, and country, except the Babuyan, Batanes,

1 at low to middle Palawan groups, in the lowlands up to 800 m groups of islands, usually et (Dickinson et al., 1991; Kennedy et al., 2000). elevations in forested areas (Dickinson al., records from The records from Camiguin consist of a female 1991; Kennedy et al., 2000). The taken from Mt. Catarman at 1500 ft (ca. 450 m) Camiguin were taken on 13 and 21 June 1968 in on 17 June 1968 (Site 9) and another female from Kasansangan at 1500-2000 ft (ca. 450-600 m) Mt. Timpoong on 13 June 1969 at an unknown and on Mt. Timpoong at 3150 ft (ca. 950 m) on elevation. Specimens of other species from Mt. 25 June 1969 (Site 13). Timpoong on the same date were taken at Specimens Examined—Total 4. Site 11 (3 3150 ft (ca. 950 m). dmnh); Site 13 (1 fmnh). Specimens Examined—Total 2. Site 9 (1 fmnh); Site 13(1 fmnh). Family Coraciidae—Rollers Eurystomus orientalis—Dollarbird —Swifts Family Apodidae The Dollarbird occurs from India to New Collocalia mearnsi— Swiftlet Philippine Guinea and the southwest Pacific, including all The Philippine Swiftlet is endemic to the of the Philippines, except the Batanes group of where it is recorded mainly on the Philippines, islands, in forest edge and clearings in the islands of Bohol, Cebu, Luzon, Mindanao, lowlands up to 1200 m (Dickinson et al., 1991; Mindoro, Negros, Palawan, and Panay, usually Kennedy et al., 2000). Dickinson et al. (1991) in forest and forest clearings above 900 m; it is noted a single specimen deposited in the Museum absent from the Babuyan, Batanes, and Sulu of Comparative Zoology, Harvard University, of islands et al., 1991; groups (Dickinson taken from Mambajao on 19 August 1921; no et from Kennedy al., 2000). Specimens Camiguin elevation was indicated. were taken from Mt. Catarman and in the Specimens Examined—None. 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 Family Alcedinidae—Kingfishers unknown elevation. Specimens of other species Ceyx lepidus—Variable Dwarf-Kingfisher from this site were taken at 3100-3350 ft (ca. The Variable Dwarf-Kingfisher occurs from 950-1000 m), with several at 4000 ft (ca. the Moluccas to New Guinea and the southwest 1200 m). Pacific islands; in the Philippines it is recorded Specimens Examined—Total 4. Site 9 (1 on the islands in the Central Philippines as well fmnh); Site 11 (1 fmnh); Site 13 (2 fmnh). as on the Mindanao and Sulu groups of islands, — in primary and secondary lowland forest (Dick- Collocalia esculenta Glossy Swiftlet inson et al, 1991; Kennedy et al., 2000). It was The Glossy Swiftlet occurs from the Anda- recorded from Mt. Timpoong at 3150 ft (ca. mans, Nicobars. and Malay Peninsula to New 950 m) on 17 and 18 June 1969 (Site 13). In May Guinea and the southwest Pacific; it is found 1994, two individuals were recorded in Kital-is, throughout the Philippines, from sea level to Sagay, at 900-1 100 m (Site 4), including one that mountaintops (Dickinson et al., 1991; Kennedy was taken in a Victor trap on 17 May. et al., 2000). It was commonly recorded on Mt. Specimens Examined—Total 4. Site 4 (2 msu-

Catarman and in the nearby area of Kasangsan- iit); Site 13 (1 dmnh, 1 fmnh).

62 FIELDIANA: ZOOLOGY Moris—White-collared Halcyon Kingfisher reference to the source, a juvenile captured in The White-collared Kingfisher occurs widely Ginsiliban in 1993. The most recent evaluation northeast Africa to from southern China, Ryu- of this species found it to be fairly common kyus, Southeast Asia to New Guinea, Australia, despite "losing ground clearly to habitat clear- and the southwest Pacific; it is found nearly all ance, hunting and trapping for trade" and over the Philippines, from exposed reefs to open assigned it a Near-Threatened status (Collar et country and forest edge (Dickinson et al., 1991; al., 1999). et It in Kennedy al., 2000). was recorded Gidag- Specimens Examined—Total 3. Site 9 (1 both in the on and Kasangsangan, vicinity of Mt. dmnh); Site 11 (1 fmnh); Site 13 (1 fmnh). Catarman, at 500-2500 ft (ca. 150-600 m; Sites 10 and 1 1) in June 1968 and on Mt. Timpoong at Family Pittidae—Pittas 3150 ft (ca. 950 m; Site 13), in Puntod at 800 ft Pitta erythrogaster—Red-bellied Pitta (ca. 250 m; Site 14), and Mantigue Island (Site The Red-bellied Pitta occurs from Sulawesi to 19) in June 1969. We recorded it further in the Moluccas and New Guinea and northeast at 10 in 1992 Balbagon, Mambajao, m May (Site Australia; it is found throughout the Philippines, 10 it 1). On May 1994, we also recorded in except the Batanes group of islands, in primary Manuyog, Sagay, at 80 m (Site 3). and secondary forest usually below 1000 m Specimens —Total 33. Site 1 Examined (1 (Dickinson et al., 1991; Kennedy et al., 2000). fmnh); Site 10 (6 dmnh, 4 fmnh); Site 11 (1 Three individuals, a male and two females, were

1 Site 13 5 dmnh, fmnh); (5 dmnh, fmnh); Site 14 taken from Mt. Timpoong at 3150-5700 ft (ca. (4 dmnh, 5 fmnh); Site 19 (1 dmnh). 950-1700 m) on 14-25 June 1969 (Sites 12 and We recorded it further on the — 13). same Family Meropidae Bee-Eaters mountain at 1475 m on 22 March 1995 (Site 8). Merops viridis—Blue-throated Bee-eater In May 1994, it was the most common bird taken The Blue-throated Bee-eater occurs in South- in Victor traps in Kital-is, Sagay, at 900-1 100 m east Asia and the Philippines, where it is known to 1200-1400 m (Sites 4-6). from many islands, except the Babuyan, Batanes, Specimens Examined—Total 9. Site 4 (2 msu- Palawan, and Sulu groups; it often found in open iit); Site 5 (1 msu-iit); Site 6 (2 msu-iit); Site 8 (1 country, scrubs, and forest clearings (Dickinson fmnh); Site 12 (1 dmnh); Site 13 (2 fmnh). et al., 1991; Kennedy et al., 2000). On Camiguin, it was recorded from Mt. Catarman and in Family Campephagidae—Cuckoo-Shrikes, Minivets, nearby Kasangsangan at 1000-4500 m (ca. 300- and Trillers 1400 m) in June 1968 (Sites 9 and 1 1) and on Mt. Lalage nigra—Pied Triller Timpoong at 3350 ft (ca. 1000 m) in June 1969 The Pied Triller ranges from the Nicobars to (Site 13). Southeast Asia; it occurs throughout the Philip-

Specimens Examined—Total 9. Site 9 (1 dmnh, pines, except the Batanes and Babuyan groups of 2 fmnh); Site 11 (2 dmnh, 2 fmnh); Site 13 (1 islands, usually in open areas in the lowlands up dmnh, 1 fmnh). to 1400 m (Dickinson et al., 1991; Kennedy et al., 2000). It was recorded from Gidag-on and Family Bucerotidae—Hornbills Kasangsangan in the vicinity of Mt. Catarman at Aceros leucocephalus—Writhed Hornbill 500-2000 ft (ca. 150-600 m) in June 1968 (Sites The Writhed Hornbill is endemic to Mind- 10 and 1 1). In June of the following year, records at 3150 ft anao, Dinagat, and Camiguin, in forest up to came mainly from Mt. Timpoong (ca. 950 Site at 800 ft 150 Site 1200 m (Dickinson et al., 1991; Kennedy et al., m; 13), Puntod, (ca. m; 2000). The records from Camiguin in 1968 14), and Mantigue Island (Site 19). consist of a male taken from Kasangsangan in Specimens Examined—Total 30. Site 10 (9 Site 11 Site 13 the vicinity of Mt. Catarman at 1000 ft (ca. dmnh, 4 fmnh); (2 fmnh); (1 Site 14 5 Site 19 300 m) on June 16 (Site 11) and another male dmnh); (4 dmnh, fmnh); (5 from Mt. Catarman at 2000-4000 ft (ca. 600- dmnh). 1200 m) on June 17 (Site 9). A female was also —Bulbuls obtained from Mt. Timpoong at 3150 ft (ca. Family Pycnonotidae — Bulbul 950 m) on 16 June 1969 (Site 13). We did not Pycnonotus goiavier Yellow-vented in record it during our fieldwork in the 1990s. The Yellow-vented Bulbul occurs widely Mallari et al. (2001) mention, without further Southeast Asia and throughout the Philippines,

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 63 of Specimens Examined—Total 94. Site 4 (8 msu- except the Babuyan and Batanes groups Site 5 Site 6 (1 msu-iit); Site 9 (6 islands, in scrub and second-growth forest iit); (2 msu-iit); dmnh, 15 fmnh); Site 10 (6 dmnh); Site 11 (9 (Dickinson et al., 1991: Kennedy et al., 2000). 7 Site 12 (2 dmnh, 2 fmnh); Site 13 It was recorded from Mt. Catarman and its dmnh, fmnh); dmnh, 13 fmnh); Site 14 (2 dmnh, 1 fmnh). nearby areas of Gidag-on and Kasangsangan at (20 500-4000 ft (ca. 150 1200 m) in June 1968 (Sites —Crows In June 1969, additional records were Family Corvidae 9-11). Crow obtained from Mt. Timpoong at 3150-4800 ft Corvus /w/oY>/7nvn/?o.v— Large-Billed Crow occurs from Iran to (ca. 950-1450 m; Sites 12 and 13) and the nearby The Large-Billed and area of Puntod at 800 ft (ca. 250 m; Site 14) as Northeast Asia, China, Taiwan, Ryukyus, it is found the well as from Mantigue Island on 28 June (Site Southeast Asia; nearly throughout in forest to and in 19). We recorded it further in Balbagon, Philippines, edge plantations et 1991; et al., Mambajao. at 10 m on 28 May 1992 (Site 1). towns (Dickinson al., Kennedy was taken from Dickinson et al. (1991) noted the presence of 2000). A male specimen Gidag- of Mt. Catarman at 1000 ft another specimen from an unspecified locality on on in the vicinity (ca. 24 June 1968 Camiguin, deposited at the Museum of Com- 300 m) on (Site 10). Examined—Total 1. Site 10 parative Zoology, Harvard University. Specimens (1 Specimens Examined—Total 30. Site 1 (1 fmnh). fmnh); Site 9 (1 dmnh, 1 fmnh); Site 10 (5 dmnh, — and Thrushes 5 fmnh); Site 11 (2 dmnh, 3 fmnh); Site 12 (1 Family Turdidae Robins,— Shamas, dmnh); Site 13 (5 fmnh); Site 14 (2 dmnh, 3 Copsychus saularis Oriental Magpie-Robin from fmnh); Site 19 (1 dmnh). The Oriental Magpie-Robin occurs Pakistan and India to southern China and Ixos everetti—Yellowish Bulbul Southeast Asia; in the Philippines, it occurs The Yellowish Bulbul is endemic to the islands throughout the country, except the Palawan, of Bucas Grande, Mind- Biliran, Dinagat, Leyte, Babuyan, and Batanes groups of islands, usually anao, Panaon, Samar, and the Sulu Siargao, in second-growth and scrubby forest (Dickinson and Camiguin, usually in primary and group, et al., 1991; Kennedy et al., 2000). It was lowland forest et al., 1991; secondary (Dickinson recorded from Mt. Catarman and in the nearby et al., 2000). Three subspecies are Kennedy area of Gidag-on at 500 3000 ft (ca. 150-900 m) including /. e. catarmanensis, which recognized, in June 1968 (Sites 9 and 10). Additional records is restricted to (Rand & Rabor, 1969). Camiguin were obtained in June 1969 from Mt. Timpoong Our of at fmnh with those comparison specimens at 4000 ft (1200 m; Site 13) and the nearby area of other named the distinc- subspecies supports of Puntod at 800 ft (ca. 250 m; Site 14). We tiveness of the Camiguin population; it is sub- recorded it further in May 1994 in Kital-is, the and darkest the three stantially largest among Sagay, at 900-1 100 m (Site 4) and 1200-1400 m races that are currently recognized (Dickinson et (Site 6). al.. 1991; Kennedy et al., 2000). The call of this Specimens Examined—Total 16. Site 4 (3 msu- species is equally distinct (Kennedy et al., 2000). iit); Site 6 (1 msu-iit); Site 9 (1 fmnh); Site 10 (4 The geographic variation observed in this species dmnh, 3 fmnh); Site 13 (3 fmnh); Site 14 (1 as defined is and is currently unusually great fmnh). suggestive of a potential species group. Further taxonomic studies are warranted. Saxicola caprata—Pied Bushchat This species was recorded in June 1968 from The Pied Bushchat occurs from Iran to Mt. Catarman and the nearby areas of Gidag-on southwest China, Southeast Asia, and New and Kasangsangan at 500-4500 ft (ca. ISO- Guinea; it is a common resident in the Philip- MOO m; Sites 9-11). Additional records were pines, except the Sulu, Palawan, Babuyan, and obtained in June 1969 from Mt. Timpoong at Batanes groups of islands, usually in fairly dry, 3150-5700 ft (ca. 950-1700 m; Sites 12 and 13) open country (Dickinson et al., 1991; Kennedy et and the nearby area of Puntod at 800 ft (ca. al., 2000). A female specimen was taken from 250 m; Site 14). In May 1994, it was one of the Mt. Catarman at 2000 ft (ca. 600 m) on 19 June most common birds mist netted in Kital-is, 1968 (Site 9).

Sagay, at 900-1 100 m to 1200-1400 m (Sites 4- Specimens Examined—Total 1. Site 9 (1 6). fmnh).

64 FIELDIANA: ZOOLOGY Zoothera andromedae—Sunda Ground-Thrush Specimens Examined—Total 31. Site 4 (1 msu- The Sunda Ground-Thrush occurs from Java iit); Site 9 (3 dmnh, 5 fmnh); Site 12 (10 dmnh, and Sumatra to the Lesser Sunda it is Islands; an 10 fmnh); Site 13 (2 dmnh). uncommon resident in the Philippines, where it has been recorded on mountains in mainly the Megalurus timoriensis—Tawny Grassbird northern and central Luzon, Mind- Mindanao, The Tawny Grassbird occurs from the Lesser oro, and in the of Negros, Panay, understory Sunda Islands, Moluccas, and Sulawesi to New forest above 1000 m (Dickinson et al., 1991; Guinea and Australia; it is found throughout Kennedy et al., 2000). We recorded a single most of the Philippines, excluding the Babuyan, specimen, taken in a Victor rat trap, from Mt. Batanes, Palawan, and Sulu groups of islands, in Timpoong at 1275 m on 23 March 1995 (Site 7); grasslands and disturbed forest up to 2000 m this is the first record for Camiguin. et (Dickinson al., 1991; Kennedy et al., 2000). It Specimens Examined—Total 1. Site 7 (1 was recorded in 1968 from Kasangsangan in the fmnh). of vicinity Mt. Catarman at 1000 ft (ca. 300 m) on June 20 — (Site 11). Additional records were Zoothera dauma Scaly Ground-Thrush obtained from Mt. Timpoong at 3250 ft (ca. The Scaly Ground-Thrush is an uncommon 980 m) in June 1969 (Site 13). We also recorded winter visitor to the Philippines from Siberia, it in May 1994 in Kital-is, Sagay, at 1000- India, China, Japan, Korea, and Taiwan. It has 1300 m (Site 5). been recorded previously only on the islands of — Specimens Examined Total 10. Site 5 (1 msu- Batan, Catanduanes, Fuga, Luzon, Marinduque, iit); Site 11 (2 fmnh); Site 13 (3 dmnh, 4 fmnh). Mindoro, Palawan, and Sibuyan, where it forages on the ground and forest understory at Cisticola exilis—Bright-capped Cisticola all elevations (Dickinson et al., 1991; Kennedy et The Bright-capped Cisticola ranges from India al., 2000). Our record on Camiguin consists of to southern China, Taiwan, Southeast Asia a single specimen taken in a Victor rat trap on (excluding the Malay Peninsula) to New Guinea, Mt. Timpoong at 1275 m on 19 March 1995 (Site Australia, and the southwest Pacific; it occurs on 7). This is the first record of this species for the most islands of the Philippines, excluding the island. Babuyan, Batanes, Palawan, and Sulu groups of — in Specimens Examined Total 1. Site 7 (1 islands, grassy habitats and rice fields (Dick- fmnh). 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 Family Sylviidae—Old World Warblers 9) and the other from Kasangsangan in the Phylloscopus trivirgatus—Mountain Leaf-War- vicinity of Mt. Catarman at 1500 ft (ca. 450 bler m) on 21 June (Site 11). The Mountain Leaf-Warbler occurs from the Specimens Examined—Total 2. Site 9 (1 Malay Peninsula, Borneo, Sumatra, Java, and dmnh); Site 11 (1 fmnh). the Lesser Sunda Islands to New Guinea; in the Philippines it is found mainly on the islands of Luzon, Mindanao, Negros, Palawan, and Panay, Family Muscicapidae—Flycatchers in montane and mossy forest above 800 m Ficedula hyperythra—Snowy-browed Flycatcher (Dickinson et al., 1991; Kennedy et al., 2000). The Snowy-browed Flycatcher ranges from An endemic subspecies, P. t. diuatae, is recog- the Himalayas to southern China, Taiwan, and nized on Camiguin and Mt. Hilong-hilong Southeast Asia; it occurs on most of the larger (eastern Mindanao; Dickinson et al., 1991; islands throughout the Philippines, in dense Kennedy et al., 2000), though Dickinson (in forest understory usually above 1000 m, but is litt.) considered this in need of reassessment. absent on Sibuyan and the Batanes and Sulu Records on Camiguin were obtained from Mt. groups of islands (Dickinson et al., 1991; Catarman at 2000^950 ft (ca. 600-1500 m) in Kennedy et al., 2000). On Camiguin, we June 1968 (Site 9) and from Mt. Timpoong at recorded this species for the first time in Kital- 4800-5700 ft (ca. 1450-1700 m) in June 1969 is, Sagay, at 900-1100 m in May 1994 (Site 4) (Sites 12 and 13). It was also recorded in Kital-is, and on Mt. Timpoong at 1275 m on 25 March Sagay, at 900-1100 m in May 1994 (Site 4). 1995 (Site 7).

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 65 from the lower Specimens Examined—Total 5. Site 4 (4 nisi 2000). It was recorded mainly of Mt. Catarman and the nearby areas of in); Site 7 (1 fmnh). slopes Gidag-on and Kasangsangan in June 1968 at ft 150-600 Sites 9 Addi- Ficedula westermanni Little Pied Flycatcher 500-2000 (ca. m; 11). records in June 1969 were taken from Mt. The Little Pied Flycatcher occurs from India tional ft 950 Site and the to southern China and Southeast Asia; in the Timpoong, at 3150 (ca. m; 13) area of Puntod at 800 ft (ca. 250 m; Site Philippines, it has been recorded only from nearby recorded it in 1994 in Kital-is, Luzon, Mindanao, Mindoro, Negros, southern 14). We also May at 1000-1400 m 5 and 6). Palawan, and Panay, in forest and forest edge Sagay, (Sites Examined—Total 32. Site 5 msu- above 800 m. usually in middle story and forest Specimens (2 Site 6 Site 9 (2 fmnh); Site 10 (7 canopy (Dickinson et al., 1991; Kennedy et al., iit); (1 msuiit); Site 1 1 3 Site 13 2000). The records from Camiguin were taken dmnh, 7 fmnh); (3 dmnh, fmnh); 14 2 from Mt. Catarman at 2000^950 ft (ca. 600- (1 dmnh, 2 fmnh); Site (2 dmnh, fmnh). 1500 m) in June 1968 (Site 9) and on Mt. in — Timpoong at 3150-5700 ft (ca. 950-1700 m) Terpsiphone cinnamomea Rufous Paradise-Fly- June 1969 (Sites 12 and 13). On 29 May 1992, we catcher recorded it on Mt. Mambajao at 1000 m (Site 2) The Rufous Paradise-Flycatcher occurs and in May 1994 in Kital-is, Sagay, at 900- throughout the Philippines, excluding the Babu- 1300 m (Sites 4 and 5). yan, Batanes, and Palawan groups of islands, in Specimens Examined—Total 42. Site 2 (1 understory of forest up to 1200 m; it also occurs fmnh); Site 4 (3 msu-iit); Site 5 (1 msu-iit); Site on several of the Talaud Islands, Indonesia 9(12 dmnh, 10 fmnh); Site 12 (9 dmnh, 4 fmnh); (Dickinson et al., 1991; Kennedy et al., 2000). Site 13 (2 fmnh). Three specimens, two females and a male, were taken on 16-23 June 1969 from Mt. Timpoong at Cyornis rufigastra—Mangrove Blue Flycatcher 1500-3150 ft (ca. 450-950 m; Site 13); another The Mangrove Blue Flycatcher ranges from female was recorded from Puntod at 800 ft (ca. the Malay Peninsula, Java, Sumatra, and Sula- 250 m) on 27 June 1969 (Site 14). In May 1994, it wesi to Borneo and the Philippines; it occurs was recorded in Kital-is, Sagay, at 900-1100 m throughout most of the islands of the Philip- (Site 4). pines, except the Batanes group, usually in Specimens Examined—Total 5. Site 4 (1 msu- disturbed forest habitat (Dickinson et al., 1991; iit); Site 13 (1 dmnh, 2 fmnh); Site 14 (1 dmnh). Kennedy et al., 2000). It was recorded from Mt. Catarman and the nearby areas of Gidag-on and Hypothymis azurea—Black-naped Monarch Kasangsangan at 500^1950 ft (ca. 150-1500 m) The Monarch ranges from India in June 1968 (Sites 9-11). Further records were Black-naped to southern China and Taiwan to Southeast obtained from Mt. Timpoong at 3150-4800 ft Asia; it occurs nearly throughout the Philippines, (ca. 950-1450 m; Sites 12 and 13) and the nearby the Batanes of islands, usually in area of Puntod at 800 ft (ca. 250 m; Site 14) in except group disturbed forest and forest edge below 1500 m June 1969. We also recorded it in Kital-is, Sagay, (Dickinson et al., 1991; Kennedy et al., 2000). at 900-1 100 m in May 1994 (Site 4). The population on is recognized as Specimens Examined—Total 70. Site 4 (4 msu- Camiguin a distinct endemic subspecies, H. a. catarmanen- iit); Site 9 (8 dmnh, 9 fmnh); Site 10 (7 dmnh, 4 sis (Rand & Rabor, 1969). Records from fmnh); Site 11 (6 dmnh, 6 fmnh); Site 12 (2 in June 1968 came from Mt. Catarman dmnh, 4 fmnh); Site 13 (7 dmnh, 5 fmnh); Site 14 Camiguin and the areas of and (4 dmnh, 4 fmnh). nearby Gidag-on Kasang- sangan at 1000^1950 ft (ca. 300-1450 m). Fur- Rhipidura javanica— Pied Fantail ther records in June 1969 came from both Mt. The Pied Fantail ranges from the Malay Catarman at 2000-4950 ft (ca. 600-1450 m; Sites Peninsula, Sumatra, and Java to Borneo and 9-11) and Mt. Timpoong at 3150 ft (ca. 950 m; the it Philippines, where occurs throughout most Site 13). In May 1994, we recorded it in Kital-is, of the islands, except the Babuyan and Batanes Sagay, at 900-1 100 m to 1200-1400 m (Sites 4- groups, usually in lowland second growth, 6). residential areas, bamboo thickets, and man- Specimens Examined—Total 33. Site 4 (2 msu- et groves (Dickinson al., 1991; Kennedy et al., 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 recorded it further in Balbagon, Mambajao, on dmnh, 3 Site 13 (12 dmnh, 2 28 1992 at 10 m (2 fmnh); fmnh). May (Site— 1). Specimens Examined Total 40. Site 1 (1 Motacillidae— Site 10 Family — Pipits, Wagtails fmnh); (15 dmnh, 21 fmnh); Site 11 (1 Anthus hodgsoni Olive Tree-Pipit dmnh); Site 14 (2 dmnh). The Olive Tree-Pipit is an uncommon migrant calvus—Coleto from eastern Asia to India, southern China, Sarcops The Coleto occurs almost Taiwan, Ryukyus, and Southeast Asia; in the exclusively in the in second Philippines it has been recorded on many of the Philippines, agricultural plantations, and forest in the larger islands, except the Batanes and Sulu growth, edge lowlands; its only record outside the is on groups, usually in pine and oak forest above country Banggi Island, off Borneo et et 300 m (Dickinson et al., 1991; Kennedy et al., (Dickinson al., 1991; Kennedy Within the 2000). On Camiguin, we recorded this species in al., 2000). Philippines itself, however, it is absent primary montane forest on Mt. Timpoong at from the Babuyan, Batanes, and Palawan of islands et 1275 m on 24 March 1995 (Site 7); it is the first groups (Dickinson al., record from Camiguin. 1991; Kennedy et al., 2000). In June 1968, two each taken Specimens Examined—Total 1. Site 7 (1 specimens were from Gidag-on at fmnh). 500-1500 ft (ca. 1 50^450 m; Site 10) and Ka- sangsangan at 1000 ft (ca. 300 m; Site 11), both — in the of Mt. Catarman. Family Artamidae Wood Swallows vicinity — Artamus leucorynchus—White-breasted Wood- Specimens Examined Total 4. Site 10 (2 Swallow dmnh); Site 1 1 (2 fmnh). The White-breasted Wood-Swallow occurs from Borneo, Sulawesi, to New Guinea, Aus- Family Nectariniidae—Spiderhunters and Sunbirds tralia, and the southwest Pacific; it occurs Anthreptes malacensis—Plain-throated Sunbird throughout the Philippines, except the Babuyan The Plain-throated Sunbird occurs throughout and Batanes groups of islands, usually at forest most of Southeast Asia; in the Philippines, it is edge or in disturbed forest up to 1800 m found on almost all islands, except the Babuyan (Dickinson et al., 1991; Kennedy et al., 2000). and Batanes groups, in coconut groves, man- Records from Camiguin in June 1968 came from groves, and second-growth forest in the lowlands Mt. Catarman and the nearby areas of Gidag-on (Dickinson et al., 1991; Kennedy et al., 2000). and Kasangsangan at 500-2000 ft (ca. ISO- Records on Camiguin came mainly from Ka- GOO m; Sites 9 and 10). Additional records in sangsangan in the vicinity of Catarman Moun- June 1969 were from Mt. Timpoong at 3150 ft tain at 1500-2000 ft (ca. 450-600 m) in June

(ca. 950 m; Site 13) and Puntod at 800 ft (ca. 1968 (Site 1 1) and Puntod at 800 ft (ca. 250 m) in 250 m; Site 14). June 1969 (Site 14). A single specimen was taken Specimens Examined—Total 19. Site 9 (1 on Mt. Timpoong at an unknown elevation on dmnh, 1 fmnh); Site 10 (5 dmnh; 4 fmnh); Site 16 June 1969. Specimens of other species from 11 (1 dmnh); Site 13 (2 dmnh; 4 fmnh); Site 14 (1 the same site and date were taken at 1500- dmnh). 3150 ft (ca. 450-950 m). — 11 — Specimens Examined Total 12. Site (3 Family Sturnidae Mynas and Starlings dmnh, 3 fmnh); Site 13 (1 dmnh); Site 14 (5 Aplonis panayensis—Asian Glossy Starling dmnh). The Asian Glossy Starling is found from Nectarinia —Olive-backed Sunbird eastern India to Southeast Asia; it occurs jugularis The Olive-backed Sunbird occurs from South- throughout the Philippines, in agricultural and east Asia to New Australia, and the residential areas and forest edge in the lowlands Guinea, southwest Pacific; in the it occurs on (Dickinson et al., 1991; Kennedy et al., 2000). Philippines, and Records from Camiguin in June 1968 came from almost all islands, except the Babuyan Batanes in disturbed the lower slopes of Mt. Catarman and nearby groups, usually heavily and below areas of Gidag-on and Kasangsangan at 500- habitats, including towns cities, et et 2000 ft (ca. 150-600 m; Sites 10 and 11). Two 1000 m (Dickinson al., 1991; Kennedy al., were obtained other specimens were obtained in Puntod at 2000). Records from Camiguin Mt. Catarman and the areas of 800 ft. (ca. 250 m) in June 1969 (Site 14). We from nearby

BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 67 Gidag-on and Kasangsangan at 500 3000 ft (ca. tional records in June 1969 were obtained from 150-900 m) in June 1968 (Sites 10 and 11). In Mt. Timpoong at 800-3150 ft (ca. 250-950 m: June 1969, additional records came from Mt. Site 13) and the nearby area of Puntod at 800 it Timpoong at 1000-5000 ft (ca. 300-1500 m; (ca. 250 m; Site 14). We obtained a further Sites 12 and 13) and the nearby area of Puntod record of it in Balbagon, Mambajao, at 10 m on at 800 ft (ca. 250 m; Site 14) as well as from the 28 May 1992 (Site 1) and in Kital-is, Sagay, at island of Mantigue (Site 19). We recorded it 900-1 100 m and 1200-1400 m in May 1994 further in Balbagon, Mambajao, on 28 May (Sites 4 and 6, respectively).

1992 at 10 m (Site 1) and in Kital-is, Sagay, at Specimens Examined—Total 63. Site 1 (1 1000-1300 m in May 1994 (Site 5). fmnh); Site 4 (6 msu-iit); Site 6 (5 msu-iit); Site

— 1 Specimens Examined Total 94. Site (2 9 (5 dmnh, 4 fmnh); Site 10 (3 dmnh, 4 fmnh); fmnh); Site 5 (1 msu-iit); Site 10 (24 dmnh, 20 Site 11 (5 dmnh, 11 fmnh); Site 13 (8 dmnh, 3 fmnh); Site 11 (4 dmnh, 6 fmnh); Site 12 (5 fmnh); Site 14 (6 dmnh, 2 fmnh). dmnh, 3 fmnh); Site 13 (1 fmnh); Site 14 (14 dmnh, 6 fmnh); Site 19 (8 dmnh). Dicaeum pygmaeum—Pygmy Flowerpecker The Pygmy Flowerpecker is common through- Nectarinia sperata—Purple-throated Sunbird out the Philippines, except Mindoro and the The Purple-throated Sunbird occurs from Babuyan, Batanes, and Sulu groups of islands, in India and Southeast Asia; it is found throughout forest, forest edge, and second growth mainly the Philippines, except the Batanes group of below 1000 m (Dickinson et al., 1991; Kennedy islands, in cultivated areas, mangroves, and et al., 2000). A single record exists from secondary forest in the lowlands (Dickinson et Camiguin, taken on 13 June 1968 in Kasangsan- al., 1991; Kennedy et al., 2000). Records from gan, in the vicinity of Mt. Catarman at an Camiguin in June 1968 were obtained from Mt. unknown elevation. A specimen of Columba Catarman and the nearby areas of Gidag-on and vitiensis was obtained on the same date and Kasangsangan at 500-3000 ft (ca. 150-900 m; from the same site at 1000 ft (ca. 300 m). Sites 9-11). In June the following year, records — Specimens Examined Total 1. Site 11 (1 were obtained from Mt. Timpoong at 3150 ft dmnh). (ca. 950 m; Site 13) and in the nearby area of Puntod at 800 ft (ca. 250 m; Site 14). Addition- Family Zosteropidae—White-eyes ally, Dickinson et al. (1991) noted two specimens Zosterops everetti—Everett's White-eye from an unspecified locality and elevation on Everett's White-eye ranges from the Malay Camiguin that are deposited at the Museum of Peninsula, Borneo, and Talaud to the Philip- Comparative Zoology, Harvard University. pines; it occurs only on the islands southeast of Specimens Examined—Total 39. Site 9 (3 Luzon, from Samar to Mindanao and Cebu to dmnh, 3 fmnh); Site 10 (6 dmnh, 6 fmnh); Site the Sulu and Tawi-tawi groups of islands, in 11 (3 dmnh, 7 fmnh); Site 13 (7 fmnh); Site 14 (1 scrub and forest to 1000 m (Dickinson et al., dmnh, 3 fmnh). up 1991; Kennedy et al., 2000). Records from Camiguin were obtained from Mt. Catarman Dicaeidae— Family Flowerpeckers and the nearby areas of Gidag-on and Kasang- Dicaeum — Flower- trigonostigma Orange-bellied sangan at 500^1950 ft (ca. 150-1500 m) in June pecker 1968 (Sites 9 11). Further records were obtained The east- Orange-bellied Flowerpecker ranges in June 1969 from Mt. Timpoong at 3150 ft (ca. ern India to Southeast it occurs Asia; throughout 950 m; Site 13) and the nearby area of Puntod at the on the and Philippines, except Babuyan 800 ft (ca. 250 m; Site 14). Batanes of islands, in forest second groups edge, Specimens Examined—Total 52. Site 9 (4 and cultivated areas below 1500 m growth, dmnh, 1 fmnh); Site 10 (11 dmnh, 9 fmnh); Site (Dickinson et al., 1991; et al., 2000). Kennedy 11 (4 dmnh, 9 fmnh); Site 13 (4 dmnh, 8 fmnh); On this is an Camiguin, species represented by Site 14 (2 fmnh). endemic subspecies, D. t. isidroi (Rand & Rabor, 1969). Records from Camiguin were obtained Zosterops nigrorum—Yellowish White-eye from Mt. Catarman and the nearby areas of The Yellowish White-eye is endemic to the Gidag-on and Kasangsangan at 500-4500 ft (ca. Philippines, where it occurs on Luzon, Mindoro. 150-1400 m) in June 1968 (Sites 9-11). Addi- Negros, and Panay and the adjacent small

FIELDIANA: ZOOLOGY islands, in forest and forest edge below 1000 m; it verified or supported by two other observers is absent from the Babuyan, Mindanao, Pala- after the two-year rule for publication. The and Sulu of islands et is wan, groups (Dickinson following the list of such species, including al., 1991; Kennedy et al., 2000). The population species identified by calls only, which when on Camiguin is the southernmost extension of verified would constitute additional records for is treated as this species' range and an endemic Camiguin and is indicative of the potentially far subspecies, Z. n. catarmanensis (Rand & Rabor, richer avifauna of Camiguin than the current 1969). Records from Camiguin in June 1968 were Species Accounts would suggest. We hope that all obtained from Mt. Catarman at 2000^1950 ft this listing will encourage other ornithologists (ca. 600-1500 m; Site 9). Additional records in and bird-watchers to conduct more studies of June 1969 were all obtained from Mt. Timpoong Camiguin birds. at 3150-5400 ft (ca. 950-1600 m; Sites 12 and 13). We recorded it further on Mt. Mambajao at Family Accipitridae 1000 m on 29 May 1992 (Site 2) and in Kital-is, Accipiter sp. Sagay, at 900-1 100 m and 1200-1400 m in May Observed by one of us (B.R.T.) along a trail 1994 (Sites 4 and 6, respectively). leading to Site 4 on 12 March 1994 between 6 am Specimens Examined—Total 181. Site 2 (2 and 3:30 pm. fmnh); Site 4 (4 msu-iit); Site 6 (3 msu-iit); Site 9 Columbidae (53 dmnh, 39 fmnh); Site 12 (36 dmnh, 15 fmnh); Family — Site 13 (20 dmnh, 9 fmnh). Ducula poliocephala Pink-necked Pigeon Observed by one of us (B.R.T.) along the same trail where an was Family Estrildidae—Avadavat, Parrotfinches, Accipiter sighted (see above), and Munias on the same date and time. Lonchura leucogastra—White-Bellied Munia Cuculidae The White-Bellied Munia ranges from the Family Cacomantis merulinus—Plaintive Cuckoo Malay Peninsula, Sumatra, Borneo, and The call of this cuckoo was noted B.R.T. at throughout the Philippines, in forest, forest edge, by Site 4 on 13 1994 as one of the most grassland, and in rice farms (Dickinson et al., May heard birdcalls at this site. The local 1991; Kennedy et al., 2000). Records of this commonly name of the bird is species on Camiguin were all obtained in Gidag- pitokai. on and Kasangsangan, near Mt. Catarman, at 500-1000 ft (ca. 150-300 m) in June 1968 (Sites Family Strigidae Otus 10 and 11). sp. The call of an unidentified scops-owl was Specimens Examined—Total 6. Site 10 (1 noted B.R.T. at Site 4 on 14 May 1994 at ca. dmnh, 3 fmnh); Site 11 (1 dmnh, 1 fmnh). by 5:40 am. Lonchura malacca—Chestnut Munia Oriolidae The Chestnut Munia is recorded from India Family and Nepal to southwestern China, Taiwan, and Oriolus sp. The call of an unidentified oriole was noted by Southeast Asia and is widespread throughout the B.R.T. at Site 4 on the same date and time that Philippines, associated mainly with rice fields, the was heard (see above). grasslands, and open country. The only records scops-owl from Camiguin were from Kasangsangan near Mt. Catarman at 1500 ft. (ca. 450 m) on 21-22 Family Pachycephalidae sp. June 1968 (Site 11). Pachycephala The call of an unidentified whistler was noted Specimens Examined—Total 3. Site 11 (1 B.R.T. at Site 4 on the same date and time dmnh, 2 fmnh). by that the scops-owl was heard (see above).

Sight Records

Following the recommendation appearing in Discussion the Rules for New Records (Kennedy et al., The bird fauna of is currently 2000), we have deliberately omitted from the Camiguin of 55 not 57 species, consisting Species Accounts all sight records that were represented by

ISLAND 69 BALETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN resident breeders and two migrants in 26 bird Mindanao, such as Basilan, Bohol, Dinagat. families. Seven species are reported here for the Leyte, Samar, or Siargao, regardless of area and first time: Ixobrychus cinnamomeus, Porzana distance from it, has a single-island endemic fusca, Rallina eurizonoides, Zoothera androme- (Dickinson et al., 1991; Kennedy et al.. 2000; dae, Zoothera dauma, Ficedula hyperythra, and Peterson et al., 2000). All these islands formed Anthus hodgsoni. Doves and pigeons (Columbi- part of the Pleistocene island of Greater Mind- dae) are the most diverse family on Camiguin, anao and thus were repeatedly connected by with eight species recorded on the island, dryland areas during much of the Pleistocene followed by flycatchers (Musicapidae) with six (Heaney, 1986, 2000; Heaney & Regalado, 1998; species, and thrushes (Turdidae) with four Steppan et al., 2003), unlike Camiguin, which species. The remaining 22 bird families are remained isolated (Heaney & Tabaranza, 2006a). represented by one to three species only. Because The current listing of birds on Camiguin is far Camiguin is a volcanic island of recent geological from comprehensive, pending more systematic origin and is separated from mainland Mind- field surveys, but based on the quite extensive anao by a sea channel that is only 10 km wide collection effort during the late 1960s, it is but 385 m deep, it would have not been worthwhile to note here the apparent absence connected to Mindanao during the many periods or depauperateness of several families that are of low sea level in the Pleistocene (Heaney, 1986; otherwise well represented on other oceanic Heaney & Tabaranza, 2006a). Thus, the current islands in the Philippines such as Sibuyan species assemblage reached Camiguin Island by (currently, the most comprehensively studied colonization, and their presence on Camiguin is small oceanic island in the Philippines for birds; documentation of the capacity of these species see Goodman et al., 1995). Sibuyan also provides for overwater dispersal. a good contrast with Camiguin in terms of the A brief comparison of the birds on Camiguin number of species that have successfully colo- with those on the closest islands, and sources of nized oceanic islands in the Philippines. Good- potential colonizers, including Mindanao and man et al. (1995) identified at least 102 resident the central Philippine islands of Negros, Panay, species on Sibuyan; Camiguin's current list pales and Cebu, is revealing. Except for the endemic in comparison. Among raptors, for instance, Camiguin Colasisi, or Hanging-Parrot, Loriculus only one eagle, S. cheela (Accipitridae), and one camiguinensis, 53 of the remaining species of owl, N. philippensis (Strigidae), have been resident birds are also found on Mindanao; recorded on Camiguin. Sibuyan, in contrast, Zosterops nigrorum does not occur on Mind- has at least six species of eagles, three species of anao. With Negros, Panay, and Cebu, Camiguin owls, and one species of falcon (Falconidae) shares 50, 47, and 42 of its resident bird species, present. A similar trend is apparent in rails respectively (Dickinson et al., 1991; Kennedy et (Rallidae), swiftlets (Apodidae), and kingfishers al., 2000). Luzon, which is farthest away from (Alcedinidae). Also noteworthy is the poor Camiguin in terms of potential source of representation of the larger doves (Columbidae) colonizers, also shares with it at least 50 resident on Camiguin that occur on Sibuyan, such as bird species (Dickinson et al., 1991; Kennedy et Ducula poliocephala, D. carola, D. aena, and al., 2000). These data indicate a remarkably high larger parrots (Psittacidae), such as Priorniturus faunal similarity with Mindanao, though it does discurus and Tanygnatlnts lucionensis (Goodman not provide clear evidence of biogeographic et al., 1995). affinity, given an equally high degree of similar- On the other hand, Camiguin and Sibuyan ity with Luzon and Panay, for instance. Thus, it appear to share the absence of species that are appears that Camiguin has been colonized by generally associated with wetlands and freshwa- widespread species. ter swamps (Anhingidae, Threskiornithidae, and The presence of at least one species endemic to Jacanidae) as well as species in high-elevation Camiguin {Loriculus camiguinensis, described in forest habitats, such as Orthotomus cucullatus, this volume) and four endemic subspecies of Bradypterus caudatus, Rhynomyias goodfellowi, birds clearly indicates that colonization rates Eumyias panayensis, Ficedula crypta, Culicicapa have been so low, even with its proximity to helianthea, Serinus estherae, and Pyrrhula leuco- Mindanao, that substantial genetic differentia- genis, which occur on some of the larger islands, tion has occurred. In this regard, it is noteworthy including Mindanao (Dickinson et al., 1991; that none of the other islands adjacent to Goodman et al., 1995; Kennedy et al., 2000).

70 FIELDIANA: ZOOLOGY Both islands also lack some forest and of general analysis the factors that influence diversity forest-edge resident birds in the following and distribution patterns of birds among families: Podargidae, Trogonidae, Capitonidae, oceanic islands in the Philippines. Chloropseidae, Paridae, and Sittidae and Lanii- Finally, we note that Camiguin is the home dae, Dicruridae, Oriolidae, Rhabdornithidae, not only to a newly described parrot (Loriculus and Timaliidae. also lacks Camiguin currently camiguinensis; Tello et al., 2006) and four records of Turnicidae, Rostratulidae, Scolopaci- endemic races of birds (Ixos everetti catarma- dae, Tytonidae, Caprimulgidae, Hemiprocnidae, nensis, Hypothymis azurea catarmanensis, Di- Picidae, Eurylaimidae, Hirundinidae, Alaudidae, ceum trigonostigma isidroi, and Zosterops ni- Oriolidae, and Pachycephalidae. grorum catarmanensis) but also to at least two It is further interesting to note that on species of mammals (Heaney & Tabaranza, Camiguin, in contrast to Sibuyan (Goodman & 2006b), one amphibian, and seven plants (Hea- Gonzales, 1990; Goodman et al., 1995) and other ney et al., 2006). It clearly warrants protection as larger and more speciose islands, several wide- a unique portion of the natural heritage of spread species associated with open country, the Philippines. We strongly recommend active cultivated areas, and disturbed habitats in the protection of the remaining forest. This will lowlands, such as Pycnonotus goiavier, Rhipidura benefit the wildlife of the island but will also be and Nectarinia are javanica, jugularis, commonly essential to the economic and social stability of found in relatively intact montane and mossy Camiguin because the mountain forests are the forest up to the peaks of Mt. Timpoong. A source of the island's essential and precious similar pattern in mammals was earlier observed water. This and other conservation issues are on Negros, which is a relatively depauperate discussed in more detail in Heaney and Tabar- island in terms of mammal diversity, where anza (2006a). 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 Acknowledgments al., 1989), and on Camiguin, where Suncus mur- inus is abundant in montane forest (Heaney & For assistance with fieldwork on Camiguin, Tabaranza, 1997; Heaney et al., 2006). The docu- we thank N. Antoque, E. Batara, N. Batocael, mentation of the above pattern of distribution N. Bojo, M. Carmona, A. DeOcampo, R. along elevational gradients has led to the hypo- Fernandez, M. Jayoma, L. Mostrales, A. T. thesis that the number of species in the native Peterson, G. Rosell, A. Tabaranza, B. Tabaranza community of small mammals in mature forest III, and D. Tabaranza. Permits were provided by determines the success of non-native small mam- the Protected Areas and Wildlife Bureau (Phi- mals on oceanic islands (Heaney et al., 1999). lippine Department of Environment and Natural Might this apply to birds on Camiguin as well? Or Resources); we especially thank A. C. Alcala, C. might habitat disturbance be an equally important Custodio, M. Mendoza, and W. Pollisco. We are factor contributing to this trend in bird diversity indebted to D. Willard for access to specimens at and distribution on Camiguin? Camiguin is the Field Museum and especially to G. Hess for a volcanically active island, with recorded erup- loans of specimens and information on collec- tions of at least two of its volcanos (Hibok-hibok tions under his care at the Delaware Museum. and Vulcan) within recent times. Mt. Hibok-hibok We wish to thank Sean Bober and Sarah Lansing erupted in 1827, 1862, 1871-1875, 1897, 1902, for assistance with preparing the manuscript. We and 1948-1958 and Vulcan in 1871 and 1874 are grateful to D. Willard and J. Bates for helpful (http://www.volcano.si.edu/gvp/world/volcano. comments on earlier drafts and to N. J. Collar cfm?vnum= 070 1-08; http://hannover.park.org/ and E. C. Dickinson for insightful reviews of the Philippines/pinatubo/page9.html). This suggests manuscript. Fieldwork was supported by the that the island has had bouts of habitat World Environment and Resources Program of disturbance brought about by these eruptions, the John D. and Catherine T. MacArthur aside from the ongoing human-induced habitat Foundation. Additional support was provided alteration and fragmentation (Heaney & Tabar- by the Marshall Field Fund, Ellen Thorne Smith anza, 2005a). Further surveys of the avifauna Fund, and Barbara Brown Fund for Mammal of Camiguin will enable a more comprehensive Research of the Field Museum.

\LETE ET AL.: ANNOTATED CHECKLIST OF THE BIRDS OF CAMIGUIN ISLAND 71 of forest mouse, Literature Cited — . 2006b. A new species genus Apomys (Mammalia. Rodentia) from Camiguin 14-27. In L. R.. Collar, N. J., N. A. Mallari, and B. R. Tabaranza. Island. Philippines, pp. Heaney. Birds of Island. Jr. 1999. Threatened Birds of the Philippines. ed.. The Mammals and Camiguin Center of Bookmark, Makati City. Philippines, a Distinctive Biodiversity. Fieldiana n.s.. 106: 1-72. Dickinson, E. C, R. S. Kennedy, and K. C. Parkes. Zoology, 1991. The birds of the Philippines: An annotated Heaney. L. R., B. R. Tabaranza. Jr.. D. S. Balete, checklist. British Ornithologists' Union Checklist and N. Rigertas. 2006. Synopsis and biogeography No. 12. of the mammals of Camiguin Island, Philippines. and Goodman, S. M., D. E. Willard, and P. C. Gonzales. pp. 28-48. In Heaney, L. R., ed.. The Mammals 1995. The birds of Sibuyan Island, Romblon Birds of Camiguin Island. Philippines, a Distinctive 106: Province, with particular reference to elevational Center of Biodiversity. Fieldiana Zoology, n.s.. distribution and biogeographic affinities. Fieldiana: 1-72. 82: 1-57. Zoology, n.s., Heideman, P. D., L. R. Heaney. R. L. Thomas, and Goodman, S. M., and P. C. Gonzales. 1990. The birds K. R. Ericsson. 1987. Patterns of faunal diversity of Mt. Isarog National Park, southern Luzon, and species abundance of non-volant mammals on Philippines, with particular reference to altitudinal Negros Island, Philippines. Journal of Mammalogy, distribution. Fieldiana: Zoology, n.s., 60: 1-39. 68: 884-888. J. B. and J. Parkin- Hauge, P., Terborgh, Winter, Kennedy, R. S.. P. C. Gonzales, E. C. Dickinson, H. son. 1986. Conservation in the priorities Philippines. C. Miranda. Jr.. and T. H. Fisher. 2000. Guide to Forktail, 2: 83-91. the Birds of the Philippines. Oxford University Heaney. L. R. 1986. Biogeography of mammals in Press, New York. Southeast Asia: Estimates of rates of colonization, Mallari, N. A. D, B. R. Tabaranza. Jr.. and M. J. extinction, and Journal of the speciation. Biological Crosby. 2001. Key Conservation Sites in the Linnean Society, 28: 127 165. Philippines. Bookmark, Makati City. — . 2000. Dynamic disequilibrium: a long-term, Ong, P., L. E. Afuang.and R. G. Rosell-Ambal. eds. large-scale perspective on the equilibrium model of 2002. Philippine Biodiversity Conservation Priori- island biogeography. Global Ecology and Biogeog- ties: A Second Iteration of the National Biodiversity raphy. 9: 59-74. Strategy and Action Plan. Philippine Department of Heaney. L. R.. D. S. Balete, E. A. Rickart. R. C. B. the Environment and Natural Resources, Quezon Utzurrum. and P. C. Gonzales. 1999. Mammalian City. diversity on Mount Isarog, a threatened center of Peterson. A. T.. L. G. Ball, and K. W. Brady. 2000. endemism on Southern Luzon Island. Philippines. Distribution of birds of the Fieldiana: Zoology, n.s., 95: 1-62. Philippines: biogeogra- and conservation Bird Conservation Heaney, L. R., P. D. Heideman, E. A. Rickart, R. B. phy priorities. International. 10: 149-167. Utzurrum, and J. S. H. Klompen. 1989. Elevational 1969. zonation of mammals in the central Philippines. Rand, A. L., and D. S. Rabor. New birds from Journal of Tropical Ecology. 5: 259-280. Camiguin South. Philippines. Fieldiana Zoology, 51: 157-168. Heaney, L. R., and J. C. Regalado, Jr. 1998. Vanishing Treasures of the Philippine Rain Forest. Steppan, S., C. Zawadski, and L. R. Heaney. 2003. A The Field Museum. Chicago. molecular assessment of phylogenetic relationships Heaney. L. R., and B. R. Tabaranza, Jr. 1997. and patterns of phylogenesis in the Philippine murid rodent Journal of Preliminary report on mammalian diversity and Apomys. Biological the Linnean 83: 699-715. conservation status in Camiguin Island. Philippines. Society, Sylvatrop(1995). 5: 57-64. Tello, J. G.. J. F. Degner. J. M. Bates, and D. E.

— . 2006a. Mammal and land bird studies on Willard. 2006. A new species of Hanging-Parrot Camiguin Island, Philippines: Background and (Aves: Psittacidae: Loriculus) from Camiguin conservation priorities, pp. 1-13. /// Heaney, L. Island. Philippines, pp. 58-72. //; Heaney. L. R., R., ed.. The Mammals and Birds of Camiguin ed.. The Mammals and Birds of Camiguin Island. a Island, Philippines, Distinctive Center of Bio- Philippines, a Distinctive Center of Biodiversity. diversity. Fieldiana Zoology, n.s., 106: 1-72. Fieldiana Zoology, n.s.. 106: 1-72.

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