Synopsis and Biogeography of the Mammals of Camiguin Island, Philippines

2006·10 MB·

by  Lawrence R Heaney

Synopsis and Biogeography of the Mammals of Camiguin
Island, Philippines
Lawrence R. Heaney,1 Bias R. Tabaranza, Jr.,1 Danilo S. Balete,1,3 and
Natalie Rigertas1
Abstract
Biodiversity surveys in the 1960s and 1990s on Camiguin Island, a geologically young,
volcanically active oceanic island surrounded by deep water, have demonstrated the presence
of24 speciesofland mammals. Three species (one insectivore and two rodents) are not native
to the Philippines, but all others (one insectivore, 12 bats, one monkey, four rodents, two
small carnivores, and one ungulate) are indigenous. Among those captured in the 1990s were
two previously unknown species ofmurid rodents in the genera Apomysand Bullimusthatare
endemictoCamiguin. Thediscoveryoftwo newspecieson suchasmall island (ca. 265 km )is
remarkable; Camiguin iscurrently thesmallest island in the Philippines known to have unique
species ofmammals. Total species richness ofnonvolant 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 notecologically balanced in the sameway as the faunas of
theislands that werepart ofGreater Mindanao: ground-livingshrews(Crocidura)and rodents
(Apomys, Bullimus, Crunomys, and Rattus) 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 onelargemammal
(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 ofsome nonvolant mammals demonstrates that dispersal across the
deep but narrow intervening channel takes place, but the presence oftwo species endemic to
Camiguin and the absence ofother species that are present on nearby Mindanao implies that
dispersal probably is rare. TheAsian house shrew (Suncusmurinus)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 ofsmall nonflying mammals was greatest at fairly high elevation.
1 Field Museum of Natural History, 1400 South Introduction
Lake Shore Drive. Chicago. IL 60605-2496. U.S.A.
2 Department of Biology, Iligan Institute of Tech- Tne Philippine Islands present a remarkable
nology, Mindanao State University, Iligan City, theaterforthestudy oftheecology andevolution
Lanao del Norte. Philippines. ofmammalian diversity. Its islands range in size
1 Laksambuhay Conservation. Inc., 10241 Mt. from less than one hectare to over 100,000 km2,
Bulusan Street, U-2. Los Banos, Laguna. Philippines. with geological age varying from under 1 million
28 FIELDIANA: ZOOLOGY, N.S., NO. 106, APRIL 5, 2006, PP. 28-48
years to over 40 million years. These islands indicated in abriefpreliminary report(Heaney&
represent many sets of historically distinct geo- Tabaranza, 1997), we found eight additional
logical units of remarkably varied origins; some species on the island that are widespread in the
had land-bridge connections to the Asian main- Philippines, plus two previously unknown en-
land in the past (those of 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 ofthe 1994and
1986, 1991b, 2000; Heaney & Rickart, 1990; 1995 mammal surveys and integrate those data
Hall, 1998, 2002). The mammals that have with information from the 1960s, including
evolved in this diverse archipelago include at information on habitat associations, relative
least 111 species endemic to the archipelago out abundance, and ecology ofthe 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
especially those surrounded by deep water (e.g., Paradoxurus hermaphroditus. Three field teams
Heaney, 1986, 2004; Goodman & Ingle, 1993; from Silliman University led by Dioscoro S.
Heaney & Tabaranza, 1997; Musseret al., 1998). Rabor collected mammals on Camiguin in 1967,
As noted by Heaney and Tabaranza (2006a), 1968, and 1969; specimens were deposited at the
Camiguin, an island of 265 km2 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),
(see Balete et al., 2006) also yielded some except as 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 otherislands 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
thatthepreviouslymeasured species richnesswas 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 alongtheelevational 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 1960s. As History (fmnh), National Museum of the Phi-
HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND 29
slliplpiepgcaiinnmeesInn(ssntmiwpte)u,treeanoald'utTMoepicsnihdenadonlaofoogrySrt(aemtpseruo-Udiniutic)vt.eirvsMeiotsiynt-- eautlnedv1ea2rt7i5torneme(Sri(otSeoitt5es),a7a)n.nddIltiinvweapvrseigmmeaotrasyttimmoonof.ntteNanonneterfaowpreperesdet
cforromwantitoon.ruTmhep slie/negtohfe(mCbRrLy)o.sSwuabsadmuelatsaunriemdalass mtaoksesnyinfoargersitcualttu1r4a7l5amrea(sSiatte185,0whe(rSeitteh3e)r,eowraisn
aredefined hereasthose that have not completed limited sampling; Table 1). This use ofhabitat is
cranial growth, especially those having unfused consistent with data from islands on Greater
basicranial sutures; these young animals have Mindanao (Rickart et al., 1993; Heaney et al.,
pelage that is usually softer and grayer than that unpubl. data).
ofadults and are noticeably lower in weight and In 1994 and 1995, three adult females were
females are usually nulliparous. Young adults trapped; one was pregnant with a single embryo
= A
are older; they have nearly completed cranial (CRL 10 mm). multiparous, nonpregnant
growth but have not yet reached adult weight female weighed 13 g; an adult male weighed 7 g.
and, usually, have not yet reproduced or are Both cranial and external measurements
pregnant for the first time. Adults have complet- (Table 2) are within stated ranges for Mindanao
ed cranial growth and adult pelage, and usually (Heaney & Ruedi, 1994) but are slightly smaller
the females are multiparous. Comments on than those ofseries taken on Biliran, Leyte, and
distribution and useofscientific names are based Maripipi (Rickart et al.—, 1993).
on Heaney et al. (1998) unless additional sources Specimens Examined Total 5. Site4 (2 fmnh,
are mentioned. Records of specimens examined 1 msu-iit); Site 5 (1 fmnh); Site 7 (1 fmnh).
are summarized at the end ofeach account; such
summaries include site number and the number Suncus murium (Linnaeus, 1766)
ofEsxpteecrinmaelnsme(ainsupraermenetnhtessesa)n.d weights reported anTdheInAdos-iAaunsthroaulsiea;-shitrenwoowccoucrcsurwsidetlhyroiunghAosuiat
here were taken in the field by members of the Philippines, though it is not native to the
the field team on fresh animals. Cranial mea- country. It is abundant in urban and agricultural
surements were taken by Heaney with digital areas; on islands with low mammal species
calipers graduated to 0.01 mm. Comparisons of richness such as Negros, it is sometimes abun-
cranial measurementsare to published records of dant in disturbed forest and occasionally in
specimens measured in the same manner by primary forest (Heideman et al., 1987; Heaney et
Heaney. al., 1989, 1991), but on islands ofaverage species
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 ofabundance is quite different from that on the
forest; and is absent outside offorest (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 ofNegros,
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 =" maximumelevation
—
1400 mossy
primary
~ J1 11. 11
1200 " montane
disturbed
— L
O 1000
_
re lowland
| 800
—
600
noforest
—
400
200
W
11 IICO </> c/> II
•oa 5) oc .£c tOQ..c0C3 OQ 5oIIccc 0) 5 ^ c"3
.5)
Fig. 1. Elevational range of nonvolant small mammals (Insectivora and Rodentia) on Camiguin Island,
Philippines. Recordsfromthe 1960sareindicatedwithstarsandfromthe 1990sbysolidsquares.Theapproximate
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
lightlydisturbed 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); Site6(10 msu-iit); Site 7 (56 fmnh); Site 8 (9
Suncus murinus was most often trapped in fmnh); Site 15 (1 dmnh).
runways orclearareasbeneath 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-
Fiveadultfemaleswithameanweightof32 ± spread in Southeast Asia and is common
4.5 g (range = 27-39 g) were pregnant; litter throughout the Philippines. It ranges from sea
m
sizes for four ofthese were one, two, three, and level to at least 1250 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
of28.8 ± 4.1 g (range = 22-35 g), and nullipa- forest (Heaney et 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
2la4r-g4e8r gt,hanNfe=mal2e7s).inMtahilsesspeacriees c(oTnasbpliec2u)o.usly ctuoramlmaorneainata1h0e0avmily(Sditiest2u;rbTeadblleow3)l.anIdt waagrsicluels-s
HEANEY ET AL.: THE MAMMALS OF CAMIGUIN ISLAND 31
Tabu 1. Numbers ofnonvolant small mammals captured in traps in heavily disturbed lowland agricultural
6a)reapr(iSmitaery3)mosenctoanndearfyorlesotwl(aSintdef7o),reastnd(Slitoewe4r).mdoisstsuyrbfeodresltow(eSirtemo8n)toanneCafmoiregsuti(nSiItsela5)n,d.prTihmearnyummboesrssyofforceasptt(uSrietse
per KM)trap-nightsaregiven in parentheses. See HeaneyandTabaranza(2005a)forfullsitedescriptions. Asterisks
mark species presumed to be present; see Methods.
r<
e a
Ctf 00
60
.£5 s>
SI es
C-o
T3 a
.;; oo
•He
o o
oo on
±j P
t"3 ac
O oo
2
c5
*
S
EQ <BuO
S c
£6
5.SE
S o.E
Q*2 a
—+i 6g §E
c a B
^Z
1
E
HIS
Tabu: 3. Numbers offruit batscaptured in mist nets in a lowland agricultural area (Site 1), heavily disturbed
lowland agricultural area (Site 3), secondary lowland forest (Site 4), disturbed lower montane forest and primary
mossy forest (Sites 5 and 6 concurrently), and primary montane forest (Site 7) on Camiguin Island during 1992,
1994,and 1995. Thenumberofcapturespernet-night aregiven inparentheses. See HeaneyandTabaranza(2006a)
for full site descriptions. Asterisks indicated species observed but not netted (see text).
Scientific name Si1t0emI, S1i5t0em3, 1S0it0e04i,n S1i2t0es0-514a0n0dm6, 1S2it7e57m,
('ynopterus brachvolis
a'S
C/3
in cranial and external measurements is slight, endemic species of Philippine pygmy fruit bats,
Camiguin specimens tend to cluster consistently Haplonycterisfischeri and Otopteropus eartilago-
within the upper ranges of most features nodus, where the phenomenon was first detected
measured (Table 4; Heaney and Rabor, 1982; (Heideman, 1989; Heideman et al., 1993; Heide-
Heaney et al., 1991, 1999, unpubl. data; Rickart man & Powell, 1998). Additionally, this condi-
et al., 1993). — tion was apparently exhibited by primiparous
Specimens Examined Total 56. Site 1 (15 young adult females only, allowing them to give
fmnh); Site 3 (1 msu-iit); Site4 (4 msu-iit); Site 5 birth only once in their first year, which had the
(8 msu-iit); Site 6 (1 msu-iit); Site 11(12 dmnh); effect of enabling them to synchronize breeding
Site 13 (5 dmnii); Site 14 (2 dmnii); Site 17 (8 with the adult females the following year (Heide-
dmnii). man & Powell, 1998).
Males are somewhat larger than females in
Ptenochirusjagori (Peters, 1861) most cranial and external dimensions, as on
The musky fruit bat is a common Philippine Biliran, Leyte, and Maripipi (Table 4; Rickart et
endemic, occurring throughout the archipelago al., 1993). Cranial and external measurements
with the exception of the Batanes/Babuyan and (Table 4) are noticeably larger than those for
Palawan faunal regions from sea level to at least series from Catanduanes and southern Luzon
1800 m (Heaneyet al., 1998). Our limited netting and were similar to those from Biliran, Leyte,
ecolplornoewimvlmCamaatanroimdyoningmfiuoon(risSensiasttyelsafohtow1or)le,w1as0etn0ldde0astsaPmg1trce3i(n0ocSo0uimclthmmteiuo4rr()anuS,lsitaaejinnra6deg;aodrTsiiacastabtrtluc1oere0b3bemi)edn. af111mn99ndS(9hp6;M)ea;cdRrimiSimncipekhtinea)psr;i4t(S(E1eiHtxt6eeaaalmmn1.s,ie3uyn-1(i,e96id9t13d)—9);m8.n4TS;hoi)ttHe;ael6aSin(t41ee6y.m1se7utS-i(iat9ile.t,)r;o11m9S)9i.1(t,8e
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
m
to at least 1200 (Fig. 2), though probably its Thailand to Australia and is found throughout
abundancedeclines with increasingelevation and the Philippineswith theexception ofthe Palawan
with increasing levels ofdisturbance (Table 3), as 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
unpubl. data; Heideman & Heaney, 1989; Rick- primary forest (Heideman & Heaney, 1989;
art et al., 1993; Lepiten, 1997). Heaney et al., 1991, 1998; Utzurrum, 1992;
Three adult females, netted in May 199=2 and Rickart et al., 1993). Records from the 1960s
1994, weighing an average of 70 g (range 68- document it from Camiguin at elevations from
(75CRg)L, w=er5e 1p0remgmn)an.tTwhirteheanosninpgrleegneamnbtryfoemeaalcehs ainbotuhte2159090tso;1b5e0c0aumse(Ftihgi.s2)s,pebcuiteswetynpeictatleldynfolniees
with large mammae had an average weight of above the canopy and our netswere set not more
73.5 g (range = 72-74 g), and two nulliparous than about 4 m above the ground, our failure to
females weighed 67 and 68.5 g. Eleven adult catch any does not necessarily indicate any
males had a mean weight of73.2 ± 4.7 g (range change in their abundance.
= 64-78 g, N = 11). Pregnant females of P. External and cranial measurements show only
j(aMgto.riIshaarvoeg)b(eeHnearneecyoredteadl.,al1s9o99i)n.MOanyMoinndLaunzaoon sDliingahgtatva,riPaatniaoyn,s Lweiytthe,tahnosdeMaorfipsippeici(mHeenasnefyro&m
(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
mlDaeaccntaetmaibnnegdrfeP(moHaweleaelnsleiy(n1e9Mt9a8a)yl..ftoouunJpnuudnbelt.haandtadtoaAn)u.gNHueesgitrdoets-o d1m7n(h3);ROMSi)t.e 12 (1 dmnh); Site 13 (2 dmnh); Site
Island, P. jagori gives birth to a single young Pteropuspwnilus Miller, 1910
twice each year: the first in late March or early The little golden-mantled flying fox is endemic
April and the second in August. It was further to the Philippines (aside from a single population
discovered that this species undergoes delayed on Miangas Island. Indonesia, adjacent to
implantation and early development that lasts Mindanao) and occurs throughout the archipel-
for five months, shorter than in two other ago, with the exception ofthe Batanes/Babuyan
36 FIELDIANA: ZOOLOGY
and Palawan regions (Heaney et al., 1998). is most often encountered from lowlands to at
Previously reported from Camiguin as P. tablasi, least 1350 m in agricultural lands to primary
the species was revised to include P. tablasi and lowland and montane forest and occasionally
P. balutus as synonyms under 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
mostcommon on smallislandsand is uncommon disturbed ha—bitats, and a larger one, designated
to rare on larger islands. Pteropuspumilus 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 (Rickartet 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
dmnh); Site 13 (2 dmnh); Site 14 (1 dmnh); Site larger than those in series from Biliran, Leyte,
17 (20 ROM). Luzon (Mt. Makiling), Maripipi, and Mindanao
— (Mt. Kitanglad), which all fall within the
Family Emballonuridae Sheath-tailed Bats dimensio—ns of the smaller morphotype, R.
Emballanura alecto (Eydoux and Gervais, 1836) arcuatus s (Ingle & Heaney, 1992; Rickart et
The Philippine sheath-tailed bat is a common al., 1993; Heaney et al., 1999, unpubl. data).
cave-dwelling species that occurs throughout Instead, theirexternalandcranialdimensionsare
most ofthe Philippines and is also known from com—parable to the larger morphotype, R. arcua-
Borneo, the Moluccas, and Sulawesi (Koopman, tus 1(Ingle& Heaney, 1992). Systematicsin this
1989).mIt hasbeen recorded only in lowland areas "species" are badly in n—eed ofdetailed study.
(450 and below) in disturbed forest and Specimens Examined Total 10. Site4 (4 msu-
agricultural areas with scattered remnant forest, iit); Site 13 (6 dmnh).
with most captures recorded from caves, under
large boulders, or in man-made tunnels (Heaney Rhinolophus inops K. Anderson, 1905
et al., 1991, 1999; Rickart et al., 1993). We did ThePhilippineforest horseshoebat iscommon
not record this species during our fieldwork in to abundant in primary lowlamnd and montane
1994 and 1995, but in 1967 specimens were taken forest from sea level to 2250 and is usually
from Tag-ibo Cave at 400 ft (ca. 120 m) and at rare in secondary forest and mossy forest
1400-3300 ft (ca. 400-1000 m) on Mt. Mamba- (Heaney et al., 1991, 1998, 1999; Rickart et al.,
jao (see also Heaney, 1984). 1993). Improvements in our understanding of
Comparison of external and cranial measure- Philippine Rhinolophus lead us to reidentify the
ments with series from Leyte and Biliran shows single specimen from Site 13, reported asWRe.
little variation (Table 4;—Rickart et al., 1993). subrufus by Heaney (1984), as R. inops.
romS)p;ecSiimteen1s8 (E2xarommi)n.ed Total 4. Site 17 (2 nfoertetsetd attwo10a0dd0itmion(Sailtema4l)e.sCirnandiiaslturabnedd elxotwelrannadl
— 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