2000

Asiatic Herpetological ResearchVol. 9, pp. 1–37

The Amphibians and Reptiles of Panay Island, Philippines
JOHN W. FERNER1, 2, RAFE M. BROWN2, 3, ROGELIO V. SISON4 AND ROBERT S. KENNEDY2
1

Department of Biology, Thomas More College, Crestview Hills, Kentucky 41017, USA. email:fernerj@thomasmore.edu; 2 Geier Collections and Research Center, Museum of Natural History and Science,
1301 Western Avenue, Cincinnati, Ohio 45203, USA; 3 Section of Integrative Biology and Texas Memorial
Museum, University of Texas, Austin, Texas 78712, USA. email: rafe@mail.utexas.edu; 4 Zoology Division,
National Museum of the Philippines, Executive House, P. Burgos Street, Manila, Philippines. email:nmzoo@fastmail.i–next.net.
Abstract.- We provide species accounts for 20 amphibians (frogs and toads) and 72 reptiles (one turtle, 36 lizards,
and 35 snakes) from the central Philippine island of Panay and its associated land-bridge islets. Panay is amoderately-sized island (123,000 km2) that currently is separated from the nearby islands of Negros, Masbate,
and Cebu by shallow seas, indicating that dry land connections once allowed free exchange of flora and fauna
between these land masses at several periods during the Pleistocene. This fact, coupled with the wealth ofknowledge on herpetological communities of Negros and Cebu, has led biologists to assume that the amphibians
and reptiles of Panay are reasonably well known. Our data suggest that this is far from true. Our recent survey
work (1989 until present) has resulted in the discovery of at least six (and probably 12–15) new species ofvertebrates, most of which appear to be high elevation rainforest Panay endemics. In this paper we note
numerous taxonomic problems that await the attention of biologists; these surely will result in an increase of the
known species from Panay. We expect that many widespread species complexes currently of uncertain taxonomic
status will eventually be recognized as Panay endemics, further bolstering the total diversity and degree ofendemism on Panay. The status of Panay herpetofauna warrants particular attention because many of thesuspected new and endemic species appear to be forest obligates. With the near complete removal of Panay’s low
elevation forests and increased pressure on its mid– and upper montane forests, we suspect that many of Panay’s
endemics are threatened by extinction. Panay should be regarded as a tropical island with a unique geologicalhistory, a spectacular compliment of habitat types, and a diverse set of species communities – all of which are
beset with a conservation crisis of deforestation and probable extinction. Panay may be a particularly appropriate
model island for large scale conservation efforts aimed at sustainable management of forest resources. However,before informed conservation and management plans can be enacted, additional biodiversity survey work must
be conducted on Panay.
Key words.- Reptiles, amphibians, SE Asia, Philippines, Panay, Visayan Islands, conservation crisis,
herpetofauna.

Introduction
The central (or “Visayan”) Philippine island of Panay(Figs. 1, 2), is part of a Pleistocene aggregate island
complex that includes Negros, Cebu, Guimaras, Masbate, Ticao, and several small land-bridge islands
associated with each of these larger land masses.
Although amphibian and reptile species similarity
among Visayan Islands was noted much earlier (Taylor, 1920, 1922a-d, 1928; Inger, 1954; Leviton,
1963c; Brown and Alcala, 1970a), it was not until
Heaney (1985, 1986) explicitly illustrated the positionof the 120 m underwater bathymetric contour in thePhilippines that the land bridge implications of channel depth became fully appreciated by biologists. It is© 2000 by Asiatic Herpetological Research

now understood that five major Philippine islandgroups, (complexes of islands separated by shallow
channels) intermittently formed much larger landmass amalgamations at various times during the midto late-Pleistocene (Fig. 1). It is presumed that these
events allowed free exchange of fauna and flora via
land-positive connections between the otherwise isolated islands of today. Each of the Philippine Pleistocene aggregate island platforms (Fig. 1) are now
recognized by biogeographers as subprovinces (Taylor, 1928) due to the fact that each supports highlycelebrated suites of endemic taxa (Taylor, 1928; Dickinson, 1991; Heaney and Regalado, 1998; Heaney etal., 1998; Alcala and Brown, 1998; Kennedy et al.,2000).

Vol. 9, p. 2

Asiatic Herpetological Research

Figure 1. The Philippines (darkly shaded islands), withthe major Pleistocene aggregate island platforms indicated by tracing of the 120 m submarine bathymetriccontour (following Heaney, 1985, 1986).

Figure 2. Panay Island, its position in the Philippines
indicated by darkened arrow (inset). Discrete shading
indicates increasing elevational increments (see key),enclosed numbers indicate collection localities (seeTable 1), underlined bold text indicates provinces(boundaries indicated with darkened dashed lines),and darkened circles indicate major cities.

2000

One unfortunate result of the prevailing perspective since Heaney’s (1985) landmark paper, has been
herpetologists’ lack of attention to islands that areland-bridge (once connected by dry land) to well surveyed members of the same Pleistocene island complex. A case in point is Panay, a central Philippine
island of 123,000 km2 with several mountain peaks ofmore than 2000 m in elevation (Figs. 1, 2). Perhaps
not unreasonably, numerous biologists over the past
century have assumed or suggested that a species’
presence on the well-surveyed Negros implied its
undocumented presence on Panay as well (see Inger,
1954; Leviton, 1963; Brown and Alcala, 1970, 1978,
1980, 1986; Brown and Rabor, 1967; Alcala, 1986).
Nevertheless, cases of low-level island endemism
have been documented on individual islands within
Pleistocene island complexes (Alcala, 1958, 1962;
Taylor, 1920; 1922a–d; 1923; 1925; Inger, 1954;
Brown and Alcala, 1961, 1970a–b, 1978, 1980,
1982a, 1986, 1994; Brown et al, 1997a–b; Brown etal., 1999), suggesting that the two large islands warrant individual attention by biogeographers. Despite
the assumption that many widespread Philippine species were present on Panay (e.g., see Alcala, 1986),
biogeographical summaries and taxonomic reviews
historically have listed only nine vouchered (based on
museum specimens) species of snakes, six geckos, six
skinks, and seven frogs (Inger, 1954; Leviton, 1963c;
Brown and Alcala, 1970, 1978, 1980) – 30% of theherpetofauna considered in the present report.
Until very recently the higher elevation forested
regions of Panay have not been explored by biologists. At present, the results of only a few preliminary
surveys in a few high elevations localities are available (Gonzales and Kennedy, 1990, 1996; Sison et al.,
1995; Ferner et al., 1997; Brown et al., 1997a; 1999).
The purpose of this paper is to synthesize museum
records that document the diversity, endemism, status,and distribution of the amphibians and reptiles fromPanay and to report on several recent collections
resulting from the National Museum of the Philippines/Cincinnati Museum of Natural History Philippine Biodiversity Inventory (1989–1994). Anotherprimary goal is to draw attention to recently-discovered endemic species of amphibians and reptiles fromPanay and to stress the need for immediate faunal surveys on this tropical island beset by a conservation
crisis of deforestation.

Material and Methods
Study sites and collection locations are presented inTable 1 and shown in Figure 2. Additional site and2000

Asiatic Herpetological ResearchVol. 9, p. 3

Table 1. Study sites and collection localities for museum specimens of amphibians and reptiles from Panay Island.
"No further data available" indicates specimens for which municipalities were the only available collection data inmuseum catalogs. The term "municipality" is synonymous with "town" but municipalities are also the politicalentity surrounding towns by these names. Thus, municipalities contain small surrounding villages (Barrios[="Barangays "] which may contain smaller "Sitios"). It is reasonable to assume that most collections localized tothe municipality level were also collected in the town of the same name. Entries including "and vicinity" indicatesspecimens that may have been collected in the municipal town, or they may be from smaller surrounding Barangays. * Combined into a single locality due to the close proximity of collection sites at the common borders ofthese municipalities.
Site

Province

Municipality

Notes

1.

Aklan

IbajayBarangay Bugtong-bato

2.

Aklan

Libacao

Libacao town "and vicinity"

3.

Aklan

MakatoMakato town "and vicinity"

4.

Aklan

Malinaono specific locality data

5.

Aklan

Malay

Nogas Point

6.

Aklan

Nabas

Nabas town "and vicinity"

7.

Aklan

Tangalan

Tangalan town "and vicinity"

8.

Aklan

Malay

Borocay Isl., no specific locality data

9.

Antique

Barbaza

Barbaza town "and vicinity"

10.

Antique

Bugasong

Bugasong town "and vicinity"

11.

Antique

Culasi

Barangay Alojipan (Mt. Madja-as;
PNM/CMNH site)

12.

Antique

Tobias Fornier

Barangay Tobias, Villaflor

13.

Antique

Valderrama

Barangay San Agustin, (Mt. Baloy andvicinity; PNM/CMNH site)

14.

Antique

San Remegio

Barangay Aningalan, Aningalan
mountain range

15.

Antique

Tibiaono specific locality data

16.

AntiqueCaluyaCaluya Isl., no specific locality data

17.

Antique

Sibay

Sibay Isl., no specific locality data

18.

AntiqueCaluyaSemirara Isl., no specific locality data

19.*

Antique

HamtikBarangay Gunisang-an

Sibalom

Barangay Egaña

San José

Barangay Bagumbayan

20.

Capiz

Capiz

no specific locality data

21.

Capiz

Marubusao

Burias22.

IloiloIloilono specific locality data

23.

IloiloIloilo City

Iloilo City "and vicinity"

24.

IloiloAjuy

no specific locality data

25.

IloiloCalinog

Mt. Baloy and vicinity

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2000

Site

Province

Municipality

Notes

26.

IloiloBarotac Viejo

Barangay San Francisco

27.

IloiloCarlesno specific locality data

28.

IloiloCarlesSicogon Island; including Buaya area

29.

IloiloDingleBulabog-Putian National Park; PNM/
CMNH Site

30.

IloiloEstanciano specific locality data

31.

IloiloLambunaono specific locality data

32.

IloiloMandurriao

no specific locality data

33.

IloiloPototanPototan town "and vicinity"

34.

IloiloCabatuanno specific locality data

35.

IloiloEstancia

Gigante North Isl., no specific locality
data

36.

IloiloEstancia

Gigante South Isl., Gabi area

37.

IloiloEstancia

Gigante South Isl., no specific locality
data

38.

IloiloCalinog

E. Catalbac ("Calinog town")

39.

IloiloEstancia

Calagna-an Isl.; Barangkalan andvicinity40.

IloiloConcepcion

ca 1 km NW SBS Iyang Beach Resort

41.

IloiloConcepcion

Pan de Azucar Isl., no specific locality
data

42.

Negros Occidental

Pulupandan

Inampulagan Isl., 8 km W Munic. W.
Pulupandan, Negros Occ. Prov.

43.

Iloilo, Subprov. Guimaras

Unknown

Guimaras Isl., no specific locality data

44.

Iloilo, Subprov. Guimaras

JordanGuimaras Isl.; Jordan town "andvicinity"45.

Iloilo, Subprov. Guimaras

Unknown

Guimaras Isl.; 2.1 km SW Daan
Banwa

46.

Iloilo, Subprov. Guimaras

Nueva Valencia

Guimaras Isl.; Nueva Valencia town
"and vicinity"

47.

Iloilo, Subprov. Guimaras

Buenavista

Guimaras Isl.; Buenavista town

48.

Iloilo, Subprov. Guimaras

Nueva Valencia

Panobolon Isl.; Nueva Valencia town.

49.

Unknown

Unknown“Masaya” (not on available maps)

50.

Unknown

Unknown

Panay Island: no Further Data

2000

Asiatic Herpetological ResearchVol. 9, p. 5

Table 2. List of known species from Panay and smaller, nearby, land-bridge islands and other islands within thepolitical boundaries or major Panay Island provinces. Entries include Panay species with vouchered specimens inmajor museum collections (see text and species accounts for discussion of taxonomy used) and the first publishedaccounts by authorities utilizing museum specimens. IR = island record or first published record from within major
Panay island provinces. 1 Previously reported from Panay, nearby islets, or Visayan sea by Alcala (1986) but without specific reference to specimens. 2 See also Gaulke (in press). 3 Recorded from land-bridge islets but currently
not recorded from the mainland of Panay. 4 A record from Semirara Isl., within the political boundary of Panay’s
Antique Province, but land-bridge to Mindoro Isl.; not likely to be discovered on Panay in the future. 5 Apparent
major range extension, in need of verification or based on specimens with locality data probably in error.

Bufo marinus

Inger, 1954

Kaloula conjuncta negrosensis

IR (see Taylor, 1920; 1922a)1

Kaloula picta

IR

Kaloula sp.

IR2

Limnonectes cf leytensis

IR

Limnonectes visayanus

Inger, 1954

Occidozyga laevis

Inger, 1954

Platymantis corrugatus

IR2

Platymantis dorsalis

IR2

Platymantis negrosensis

Sison et al., 1995

Platymantis insulatus

Brown and Alcala, 1970b3

Platymantis panayensis

Taylor, 1920; Inger, 1954; Brown et al., 1997a

Platymantis sp. 1

IR

Platymantis sp. 2

IR

Platymantis sp. 3

IRRana cancrivora cancrivora

Inger, 1954

Rana erythraea

Inger, 1954

Rana cf. everettiSison et al., 1995

Rana vittigera

IR

Polypedates leucomystax

Inger, 1954

Cuora amboinensis amboinensis

Gaulke and Fritz, 1998

Bronchocela sp.

Taylor, 1922cDraco spilopterus

Musters, 1983

Hydrosaurus pustulatus

IR2

Gonocephalus sp.

Sison et al., 1995

Cosymbotus platyurus

Brown and Alcala, 1978

Cyrtodactylus annulatus

IR2Vol. 9, p. 6

Asiatic Herpetological Research

Cyrtodactylus philippinicus

IR

Gehyra mutilata

Brown and Alcala, 1978

Gekko gecko

Taylor, 1922c; Brown and Alcala, 1978

Gekko gigante

Brown and Alcala, 19783

Gekko mindorensis

Sison et al., 1995

Hemidactylus frenatus

Brown and Alcala, 1978

Hemidactylus stejnegeri

Sison et al., 1995

Hemiphyllodactylus insularis

Brown and Alcala, 1978

Lepidodactylus lugubris

Brown and Alcala, 19783

Lepidodactylus planicaudus

Brown and Alcala, 1978

Brachymeles boulengeri taylori

IR

Brachymeles talinis

Brown and Alcala, 1980

Brachymeles tridactylus

Brown and Alcala, 1980

Dasia grisea

Brown and Alcala, 19804

Dasia semicincta

IR5

Emoia atrocostata

IR3

Lamprolepis smaragdina philippinica

Brown and Alcala, 1980

Lipinia pulchella taylori

IR

Mabuya indeprensa

Sison et al., 1995

Mabuya multicarinata borealis

IR

Mabuya multifasciata

IR2

Parvoscincus sisoni

Ferner et al., 1997

Sphenomorphus arborens

Brown and Alcala, 1980

Sphenomorphus coxi divergens

IR5

Sphenomorphus cumingi

IR5

Sphenomorphus jagori grandis

Brown and Alcala, 1980

Sphenomorphus steerei

Brown and Alcala, 1980

Tropidophorus grayi

Sison et al., 1995

Varanus salvator nuchalis

Gaulke, 1991a, 1991b, 1992

Acrochordus granulatus

IR

Python reticulatus

Leviton, 1963c

Ahaetulla prasina preocularis

Leviton, 1963c, 1968

Boiga angulata

IR2

2000

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Asiatic Herpetological ResearchVol. 9, p. 7

Boiga cf cynodon

IR2

Boiga cf. dendrophilaIR

Calamaria gervaisi

Leviton, 1963c; Inger and Marx, 1965

Cerberus rynchops

Gyi, 1970

Chrysopelea paradisi

Sison et al., 1995

Cyclocorus lineatus alcalaiIR

Dendrelaphis caudolineatus terrificus

Leviton, 1970b

Dendrelaphis pictus pictus

Leviton, 1963c, 1970b

Elaphe erythrura psephenoura

Leviton, 1979

Gonyosoma oxycephala

IR2

Hologerrhum dermali

IR2

Lycodon aulicus capucinus

Leviton, 1965

Oligodon modestum

Sisoni et al., 1995

Psammodynastes pulverulentus

IR2

Pseudorabdion mcnamarae

Sisoni et al., 1995

Pseudorabdion oxycephalum

IR

Pseudorabdion talonuran

Brown et al., 1999

Tropidonophis negrosensis

Leviton, 1963c; Malnate and Underwood, 1988

Zaocys luzonensis

Ross et al., 1987

Calliophis calligaster gemianulis

Leviton 1963b, 1963c

Hydrophis belcheri

IR1

Hydrophis cyanocinctus

IR1

Hydrophis elegans

IR

Hydrophis inornatus

IR

Lapemis hardwickii

IR1

Laticauda colubrina

IR

Ramphotyphlops braminus

IR

Rhamphotyphlops cumingii

IR

Typhlops castanotus

Wynn and Leviton, 1993

Typhlops hypogius (= T. ruber ?)IR? (see McDiarmid et al., 1999)

Tropidolaemus cf wagleri

IR

Trimereserus flavomaculatus

Gaulke (in press)

Vol. 9, p. 8

Asiatic Herpetological Research

habitat information for many TNHC, PNM, CAS,
FMNH, and CMNH specimens may be found in themuseum catalogs. The PNM/CMNH Philippine
Biodiversity Inventory team conducted field studies atsites 11 and 13 (Figs. 3–9) and various members of
the team (particularly RVS and RMB), visited otherlocations (Figs. 10–14) to do more limited surveying.
We established elevational transects in a variety ofhabitat types (Ruedas et al., 1994, as modified byBrown et al. 1995a, 1996, in press) and utilized standard collection and specimen preservation techniques(Simmons, 1987; Heyer et al., 1994). Detailed examination of all material was conducted by RMB andJWF.
We follow the taxonomy of Taylor (1922c), Brown
and Alcala (1978, 1980) and Brown et al. (1995a;
1995b) for gekkonid and scincid lizards. The taxonomy of Inger (1954, 1996), Frost (1985), Duellman(1993), Brown and Alcala (1994), Inger and Tan(1996), Alcala and Brown (1998) and Emerson et al.
(2000), was consulted for amphibians and we include(in parentheses) the unsubstantiated taxonomic
hypotheses of Dubois (1992) for reference. While nosuitable taxonomy currently is available for Philippine
agamids (see Taylor, 1922c), we consulted Musters(1983) and McGuire and Alcala (2000) for identification of our Draco specimens. Snake taxonomy was
based on Taylor (1922a), Leviton's “Contribution to a
Review of Philippine Snakes” series (Leviton 1985,
1961, 1962, 1963a–c, 1964a–d, 1965, 1967, 1968,
1970a–b, 1979, 1983), Leviton and Brown (1959),
Inger and Marx (1965), Malnate and Underwood(1988), Wynn and Leviton (1993), McDiarmid et al.
(1999), and Brown et al. (1999). Scale counts on
snakes were taken using the methods of Dowling(1951a, 1951b). Field techniques in our surveys havebeen described in Ruedas et al. (1994) and Brown etal. (1995a; 1996; in press). Museum abbreviations follow Duellman et al. (1978) and Leviton et al. (1985).
In order to examine large scale faunal similarities
between Panay and other large island of the Philippines, we assessed overall herpetofaunal simmilarities
by calculating coefficients of similarity using a simple
index (C = 2W/a+b; see Brown and Lomolino, 1998,
for review) for the amphibians and reptiles of Panayand other islands. Our data for these calcualteions
were based on all available literature (above) andupdated conservative estimates of the taxonomy ofPhilippine amphibians and reptiles (Brown, Crombie,
Diesmos, unpublished data).

2000

Results
We present records for 20 amphibians (anurans; Figs.
15–25), and 72 reptiles (one turtle, 36 lizards, and 35
snakes; Figs 26–52) from Panay and its associatedland-bridge islets. The results of faunal similarity calculations are presented in Figure 53. Individual species accounts, with comments on their status, arepresented below.

Species Accounts
Amphibia
Anura
Bufonidae
Bufo marinus (Linnaeus) (Fig. 15)
Introduced on most inhabited islands of the Philippines (Inger, 1954; Diesmos, 1998; Alcala and Brown,1998), this species is particularly common in agricultural areas and near human habitation. It is widespread and abundant in disturbed and agriculturalareas on Panay.
Localities and specimens: (Site 1) PNM 1144–47,
1228–33; (Site 11) CMNH 4958–59, PNM 1289–96,
2552–54; (Site 12) PNM 1232–33; (Site 19) CMNH
5197; (Site 23) USNM 339985–86; (Site 24) FMNH
61482–84; (Site 31) PNM 1144–47; (Site 33) USNM
339987–88, 340062.

Microhylidae
Kaloula conjuncta negrosensis Taylor (Fig.
16)
Taylor (1920, 1922a) first collected this form on
Negros and Guimaras islands. He (Taylor, 1922a) recognized it as a full species but Inger (1954) placed K.
negrosensis in the synonymy of Luzon’s Kaloula conjuncta; no further taxonomic studies have been forthcoming. Kaloula conjuncta negrosensis is both aburrower and a climber (Taylor, 1920; Alcala, 1962,1986); it has been found in the detritus of the forest
floor (750–1075 m on Negros; Alcala, 1962) and in
the leaf axils and holes of trees (Alcala 1962). They
also emerge and congregate around water that collects
in pools and ditches in the rainy season (Taylor, 1920,
1922a). Our Mt. Madja-as specimen was found in atree hole 10 cm from the ground (tree < 0.5 m dbh).
Specimens from near the base of Mt. Baloy were
found in breeding aggregations around water buffalo
wallows near a large river. We suspect that the2000

Asiatic Herpetological Research

Visayan populations represent a distinct evolutionary
lineage and that they will eventually once again berecognized as a full species once new data become
available (Brown and Diesmos, unpublished data).
Localities and specimens: (Site 8) CAS 127890;(Site 11) PNM 2555; (Site 13) TNHC 56340–46; (Site16) CAS 127510–11, 127591; (Site 18) CAS 127815;(Site 47) CAS 124446.

Kaloula picta (Dumeril and Bibron) (Fig. 17)
This frog is found at low elevations (100–200 m) inopen areas near human habitations (Boulenger, 1882;
Alcala, 1956, 1958). It is a burrowing species thatconceals it self under leaf litter and top soil until the
beginning of the rainy season. Choruses may containhundreds of individuals (Alcala, 1962; Alcala and
Brown, 1998). This appears to be the first publishedaccount of this species on Panay.
Localities and specimens: . (Site 16) CAS 127617–
37; (Site 18) CAS 127816, 127827; (Site 23) CAS–
SU 14219–20; USNM 78079–80, 78842.

Kaloula sp.
In 1992, while conducting survey work at Barangay
Alojipan (Site 11), we heard the distinctive honking
calls of a forest species of Kaloula. Although we wereunable to locate and collect specimens, we noted thatthe calls were superficially similar to Kaloula kalingensis (Taylor, 1922a; single honk, delivered approximately every 15 to 20 minutes) from Luzon. Neither
Kaloula kalingensis nor the related K. kokacii (Rossand Gonzales, 1991) have been reported from the
nearby island of Visayas, although recent field work in
the last remaining low elevation forests of Negros(Municipality of Ayungon, Negros Oriental Prov.; C.
N. Dolino and A. C. Diesmos, personal communication) reveals the presence of a forest species (also previously unrecorded) with single honking calls there as
well. Gaulke (in press) recently has discovered a population of Kaloula in NW Panay that may be the sameas that heard (but not collected) previously on Panay
and Negros.
Localities and specimens: none.

RanidaeLimnonectes (= “Limnonectes”) cf. leytensis(Boettger) (Fig. 18)
This species is widely distributed in patches and considered common in some localities on the nearby
Negros (Alcala, 1986; Alcala and Brown, 1998).
However, it has been collected from only one locality

Vol. 9, p. 9

on Panay (Sison et al., 1995). On Negros, this species
inhabits coolor high elevation mountain streams
between 150 and 900 m (Alcala, 1962) and probably
breeds terrestrially but deposits eggs in close proximity of water (Alcala and Brown, 1956; Alcala, 1962;
Brown and Alcala, 1982b). When hatching, terrestrial
embryos of these and related species fall, are carriedby males, or are washed into water (Alcala, 1962; seealso Inger et al., 1986; Brown and Iskandar, 2000).
We find it doubtful that Visayan specimens identified
as Rana cf leytensis are conspecific with specimens
referred to this species from the Mindanao Aggregate
Island Complex (Leyte, Samar, Bohol, and Mindanao).
Locality and specimens: . (Site 25) PNM 1114–15.

Limnonectes (= “Limnonectes”) visayanus(Inger) (Fig. 19)
This large fanged frog is found in clear forest streams;
it is known to hide in rock crevices during the day andcall from rocks and stream banks above water at night(Alcala, 1962; Alcala and Brown, 1998). We found L.
visayanus at low elevations on rocks in large rivers
near Mt. Madja–as. This species may breed and lay
eggs outside of water (Alcala, 1962).
Localities and specimens: (Site 2) PNM 1715–20;(Site 3) CAS 137592–95, 137590–91, 139164–66,
USNM 305671–76; (Site 4) PNM 1613–20, 1623–27;(Site 6) CAS 137596–98; (Site 7) PNM 1799–800,
1828–31, 1836–39, 1845, 1855–60, 1865–77; (Site11) CMNH 4894–98, 4899, PNM 1302–06, 2617–21;(Site 13) TNHC 56337; (Site 14) PNM 3710–12,
3732, 3764–68, 3805; (Site 25) 1085–92, 1133–38,
1140; (Site 28) CAS 124093–106, 124442–44,
124950–58; (Site 39) CAS 124121, 124293–97; (Site44) CAS 125308–309, 125312; (Site 47) CAS
125302–307; (Site 50) USNM 78072–78.

Occidozyga laevis (Günther) (Fig. 20)
Occidozyga laevis is found in flooded fields in agricultural areas, in road–side ditches and open sewers,
and in streams and rivers from lowlands to high elevation forested sites (Inger, 1954; Alcala, 1962; pers.
obs.). On Negros this species has an altitudnal rangeof sea level to 1150 m (Alcala, 1962). Specimens arecommon in stream-side pools along larger rivers inforested areas near Mt. Madja-as and Mt. Baloy.
Localities and specimens: . (Site 1) PNM 1110–11,
1113, 1116–29, 1141, 1184–99; (Site 2) PNM 1116–
29, 1690–1714, 1721–23, 1731, 1757–58; (Site 3)
CAS 137586–88, 139148, 139167–68, USNM
305647–48; (Site 4) PNM 1600–02, 1611, 1621–22;

Vol. 9, p. 10

Asiatic Herpetological Research(Site 6) CAS 137614–15, USNM 305649, 305650–
54; (Site 7) PNM 1832–35, 1841–44, 1848–54, 1862–
64, 1880; (Site 9) PNM 1156, 1165–68, 1170–71;(Site 10) PNM 1110–11, 1113; (Site 11) CMNH
4951–57, PNM 1329, 2655–61; (Site 12) PNM 1163–
64; (Site 14) PNM 3730–31, 3782, 3804; (Site 22)
CAS–SU 9813; (Site 23) CAS–SU 14224–25, 14373;(Site 24) FMNH 61478–81; (Site 25) PNM 1141;(Site 26) PNM 1160–62; (Site 27) CAS–SU 14049;(Site 28) CAS 124959–70, 124059–76, 124426,
124432–33, 124439; (Site 30) CAS–SU 14223; (Site32) PNM 1172–73; (Site 38) CAS 132880, 132887–
901, 134089–96; (Site 39) CAS 124171, 124190–91;(Site 40) USNM 339989; (Site 41) CAS 125001,
124177, 124184–85, 124194–96; (Site 42) CAS–SU
23946–49, 23952–58, 23961–63; (Site 44) CAS
125361–62; (Site 47) CAS 125311, 125344–59.

Platymantis corrugatus (Dumeril) (Fig. 21)
This widely distributed terrestrial frog inhabits theforest floor from sea level to above 1300 m (Alcala,1986). On Mt. Madja-as we found P. corrugatus inleaf litter and in limestone crevices. This account andthat of Gaulke (in press) appear to be the first published records of this species from Panay.
Localities and specimens: (Site 6) CAS 137616–19,
139149, 185494; (Site 11) CMNH 4960–63, 5118,
PNM 2556–59; (Site 13) CMNH 3160–65; (Site 25)
PNM 1103–06; (Site 28) CAS 124058.

Platymantis dorsalis (Dumeril)This common forest frog is found in the detritus of theforest floor as well as in tree cavities and low tree
ferns (Alcala, 1962; Alcala and Brown, 1998).
Although recent studies (Brown et al., 1999) indicate
the presence of numerous cryptic species in the P.
dorsalis complex on Luzon (and we suspect that further new species await discovery in the Visayan
islands), the calls of some P. dorsalis have been heard
on Mt. Baloy and Mt. Madja-as are, at least superficially, similar to the short, whistling, ascending frequency sweep of true P. dorsalis from Luzon Island(Brown et al., 1997c). Thus, while we expect that
more species in the dorsalis complex will soon be discovered in the Visayas, we can confidently assert thatat least some Panay populations are indistinguishable
from P. dorsalis of Luzon Island (Brown et al., 1997c;
1999). This account and that of Gaulke (in press)
appear to be the first published records of this species
from Panay.
Localities and specimens: . (Site 2) PNM 1734–56;(Site 6) CAS 137620–40, 137649–51, 139150–63,
USNM 305655–70; (Site 11) CMNH 4964–98, 5206,

2000

PNM 2559, 2562–88; (Site 13) TNHC 56347–50;(Site 14) PNM 3713–19, 3729, 3733–45, 3756–62,
3772–74, 3783–85, 3788–89, 3791–96, 3830–31,
3857, 3860, 3862–65, 3882–83, 3886–88, 3893–95,
3906–08; (Site 25) PNM 1093–102; (Site 28) CAS
124419, 124428–31, 124440–41, 124689–91; (Site39) CAS 124123–33, 124146–47; (Site 41) CAS
124041–42, 124122, 124186–89, 125014–19.

Platymantis negrosensis Brown, Alcala,
Alcala, and Diesmos
This recently-described forest frog (Brown et al,
1997b) has only been documented from two sites on
Panay but is also known from localities on the nearby
island of Negros, from approximately 300 to 1625 m
in elevation (Alcala, 1958; Alcala and Brown, 1957;
Alcala, 1962). On that island, this species occupies
arboreal microhabitats in primary forest (Brown andAlcala, 1961; Alcala, 1962; Brown et al, 1997b). We
documented this island record for Panay (Sison et al.,
1995; then recognized as P. guentheri) from specimens collected in forest on Mt. Baloy at about 950 m.
This species is related to P. luzonensis but differs incharacteristics of the advertisement call an external
morphology (Alcala and Brown, 1998).
Localities and specimens: . (Site 13) CMNH 3166;(Site 14) PNM 3889.

Platymantis insulatus Brown and Alcala (Fig.
22)
A frog known from primary and secondary forest situated on karst limestone outcrops, this species was
originally discovered (Brown and Alcala 1970b) on
the forest floor and in the open mouths of small caves
at low elevations (Alcala and Brown, 1998) on theisland of Gigante South off Panay's northeast coast. A
recent (June 2000) visit to the type locality by R.
Brown and A. Alcala confirmed this species’ persistence despite the complete removal of the original forest. The presence of an endemic species on such asmall, land-bridge island is puzzling and suggests thatit may still be (or at least, may have once been)present on eastern Panay. Unsurveyed limestone formations along the northeastern coast are the mostpromising possibility for locating this species on
Panay.
Localities and specimens: . (Site 35) CAS 157235–
39; (Site 36) CAS 137641–42; (Site 37) CAS
117440–41; 119967–69, MCZ A-72946.

2000

Asiatic Herpetological Research

Platymantis panayensis Brown, Brown andAlcalaOnly recently described (Brown et al., 1997a) fromour collections from Mt. Madja-as, this species isclosely related to P. hazelae from Negros and occupies similar microhabitats in high elevation cloud forests. Taylor (1920:101) apparently had a specimen(collected by R. McGregor) of this species on handduring the description of Philautus (= Platymantis)
hazelae and he considered it conspecific with the
Negros population. On the basis of Taylor’s (1920)
account, Inger also (1954) included P. hazelae in his
list of species from Panay .
We collected the majority of the type specimens of
this species from leaf axils, leaves on shrubs, and theleaf litter on the forest floor. The call has not yet beenformally described, but consists of a pure, ringing,
tonal note with no frequency or amplitude modulation; it sounds to the human ear like the sound produced by the ringing of a small bell (1–2 notes/s).
Localities and specimens: (Site 6) CAS 137641–42;(Site 11) PNM 2314–18, 2495, 2589–90; CMNH
4113–15, 4116–20, 4868–69.

Platymantis sp. 1

Several immature specimens of a tiny, darkbrown, tuberculate (11–15 mm SVL) speciesof Platymantis were taken at high elevations
on Mt. Madja-as and on Mt. Baloy. They
appear most similar to Platymantis pygmaeus of the Sierra Madre mountains ofLuzon's east coast. Due to the fact that theavailable specimens are all sexually immature, we cannot recognize them taxonomically until further material and recordings ofadvertisements calls become available.
Localities and specimens: (Site 11) CMNH 8132
(Site 13) CMNH 3173–74, 3177.

Platymantis sp. 2
Two immature specimens of a diminutive, black,smooth-skinned (12–13 mm SVL) species of Platymantis were collected at 1450 m from under leaf litter.
The small sample size and immaturity of the specimens necessitates that taxonomic recognition of thespecies must await the collection of further material.
Locality and specimens: (Site 11) CMNH 8133–34.

Platymantis sp. 3This unidentified species is represented by a single,
very large, black specimen with two dorsolateral lightVol. 9, p. 11

lines; it is appears possibly related to P. pseudodorsalis from Luzon (Brown et al., 1999).
Locality and specimen: (Site 6) CAS 185495.

Rana (= “Fejervarya”) cancrivora cancrivora
Gravenhorst
This common frog is found in swamps, ponds,
flooded rice fields and ditches (Inger, 1954). It isfound in almost any pool of water at low elevations(Alcala and Brown, 1998). Inger (1954) first reported
it on Panay Island.
Localities and specimens: . (Site 1) PNM 1178–82;(Site 8) CAS 127893–95, 127899–904; (Site 9) PNM
1156; (Site 16) CAS 127509, 127559–81, 127611,
127678–79; (Site 17) CAS 127800; (Site 18) CAS
127801–803; (Site 19) PNM 2607–16; (Site 22)
USNM 78862–902; (Site 23) CAS–SU 14259–60,
14452–79, 15486–514, USNM 77984–96, 77988–
78042, 78048, 78062–65; (Site 24) FMNH 61398–
402; (Site 26) PNM 1160–62; (Site 27) CAS–SU
14513–25; (Site 28) CAS 124262–71, 185665–69;(Site 30) CAS–SU 14252–57; (Site 34) CAS–SU
9762–63; (Site 36) CAS 124343; (Site 37) CAS
124564–76; (Site 38) CAS 132878–79; (Site 39) CAS
124135, 124175; (Site 40) USNM 339990, 340059–
60; (Site 41) CAS 124178–79, 124320–28, 125020–
28; (Site 42) CAS–SU 23966; (Site 44) CAS 125194–
96; (Site 46) CAS 125183–93; (Site 47) CAS
125174–82; (Site 48) CAS 124791–92; (Site 50)
USNM 78066–71.

Rana (= “Hylarana”) erythraea (Schlegel) (Fig.
23)
This widely-distributed and common frog is believed
to have been introduced to the Philippines (Diesmos,
1998; Alcala and Brown, 1998), originally on Negros(Inger, 1954; Alcala, 1962; Alcala and Brown, 1998).
It's presence on Panay has been previously documented (Taylor, 1920; Inger, 1954). We found our
specimens along the grassy boarders of flooded rice
fields.
Localities and specimens: (Site 2) PNM 1725–26;(Site 3) CAS 137589; (Site 4) PNM 1603–10, 1612;(Site 7) PNM 1840; (Site 8) CAS 127891–92,
127905–912; (Site 10) PNM 1174–76; (Site 11)
CMNH 4870, PNM 1309–28, 2591–92; (Site 14)
PNM 3816; (Site 19) PNM 2622–54; (Site 22) CAS–
SU 9744; (Site 23) CAS–SU 14537–47, USNM
77730–983, 78043–47, 78049–61; (Site 24) FMNH
61391–97, 61449—52; (Site 27) CAS–SU 14578–84;(Site 30) CAS–SU 11120–24, 14526–36, FMNH
40527; (Site 33) USNM 38650–54; (Site 38) CAS

Vol. 9, p. 12

Asiatic Herpetological Research

132881–86, 134086–88; (Site 39) CAS 124209–212,
124214; (Site 44) CAS 125158–69; (Site 47) CAS
124143–57, 125310; (Site 50) USNM 77617–729,
78413–38.

Rana (= “Chalcorana”) cf. everetti Boulenger
Species of the Rana everetti complex are found in andalong streams from 300 to about 1300 m (Inger, 1954;
Brown and Alcala, 1955; Alcala, 1962; Alcala, 1986;
Brown et al., 2000). Our Mt. Baloy expedition in 1989
first recorded the presence of this frog on Panay(Sison et al., 1995). Negros populations (expected tobe conspecific with those on Panay) are only foundnear water during breeding; this species has mostoften been collected in overhanging, streamside vegetation (Brown and Alcala, 1955; Alcala, 1967). As
noted by Brown et al. (2000), the taxonomic status of
the Visayan populations referred to Rana everetti is inneed of further study.
Localities and specimens: . (Site 2) PNM 1732–33;(Site 14) PNM 3771, 3800–03, 3806–14, 3817–24,
3896, 3913; (Site 39) CAS 124213, 124215–16.

Rana (= “Fejervarya”) vittigera Wiegmann(Fig. 24)
This species occurs in open, agricultural areas nearsources of water (ponds, flooded rice fields). The distinctive, rapid honking call of this species can beheard in choruses of up to hundreds of individuals.
This record appears to be the first published accountof this species from Panay.
Localities and specimens: . (Site 11) CMNH 4871–
72, PNM 2593–94; (Site 19) PNM 2596–606; (Site40) USNM 339991; (Site 41) CAS 124197; (Site 46)
CAS 125360.

Family Rhacophoridae
Polypedates leucomystax (Gravenhorst) (Fig.
25)
This common tree frog is widely distributed in Panay(Inger, 1954; Alcala, 1986) in agricultural areas, forest edges, and disturbed forests. It ranges from nearsea level to 1000 m on Negros (Alcala. 1962) and was
first documented on Panay by Inger (1954). Our specimens were collected in banana plantations and rice
fields near the base of Mt. Madja-as and Mt. Baloy.
Localities and specimens: . (Site 1) PNM 1107–09,
1209–27; (Site 2) PNM 1729–30; (Site 6) CAS
137599; (Site 7) PNM 1797–98; (Site 11) CMNH
4997–98, PNM 2560–61; (Site 13) TNHC 56336,
56338; (Site 14) PNM 3763, 3799; (Site 16) CAS

2000

127512–14, 127540–58, 127592–95, 127608–10,
127647–49, 127657, 127670–76, 127680–97,
127701; (Site 17) CAS 127721–26; (Site 18) CAS
127841; (Site 24) FMNH 61485; (Site 25) PNM 1107–
09; (Site 28) CAS 124110–13, 124420, 124422–25,
124581; (Site 29) USNM 339992–93; (Site 30) CAS–
SU 11113–16, 14764, FMNH 40569, 44263; (Site 37)
CAS 124114, 125034–37; (Site 39) CAS 124158,
124192, 124204–07; (Site 40) USNM 339994; (Site41) CAS 124176, 125031–32; (Site 42) CAS–SU
23950–51, 23959–60, 23964–65; (Site 47) CAS
125342–43.

Reptilia
Testudines
Bataguridae
Cuora amboinensis (Daudin) (Fig. 26)
This common species is found throughout the Philippines (Gaulke and Fritz. 1998) and on Panay (Gaulke,in press) in low elevation forests, agricultural areas,
and near streams and swampy areas; it is generally
considered nocturnal (Alcala, 1986). We have collected this species in stream-side habitats at low elevations on Panay.
Localities and specimens: (Site 7) PNM 1888–89,
1891–95; (Site 11) CMNH 5500, 5501, 5502; PNM
1288, 5657–58; (Site 23) USNM 78103–04, 78746–
49; (Site 41) CAS 153872; (Site 45) CAS 185507;(Site 50) USNM 78081–102.

Squamata (Lizards)
AgamidaeBronchocela cristatella (Kuhl) and B. marmorata (Gray)
These arboreal lizards (Alcala, 1986) are found fromlowland cultivated areas to lower midmontane primary and secondary forests; they are most often
encountered sleeping at night in stream-side vegetation. No suitable taxonomic reference is available forPhilippine Bronchocela and so the true identities of
Panay specimens referred to B. cristatella and B. marmorata are uncertain. There is little consensus regarding the identity of Bronchocela throughout the
Visayas. Although specimens from Negros and Panay
key out to earlier descriptions of both Bronchocela
marmorata and B. cristatella (Taylor, 1922c; Alcala,1986), both “species” appear to be highly variable anddiagnostic characters vary ontogenetically. We con-

2000

Asiatic Herpetological Researchsider it unlikely that two independent lineages occurin sympatry on Panay and, at present, we hold in
abeyance the identity of these populations until a thorough taxonomic revision is available.
Localities and specimens: . (Site 6) CAS 137605;(Site 22) CAS–SU 10948; (Site 23) USNM 77133–
38, 78105–107; (Site 41) CAS 124333; (Site 47) CAS
125337–38.

Draco spilopterus (Weigmann) (Figs 27, 28)This species is common at lower to mid-montane elevations and often is found in coconut groves and forest edges (Alcala, 1986; McGuire and Alcala, 2000).
Draco spilopterus is the only species currently recognized from the Visayan and Luzon aggregate island
complexes, despite biogeographic and morphologicalevidence suggesting that Luzon and Visayan populations constitute independent evolutionary lineages(Taylor, 1922c; Heaney, 1985, 1986; McGuire andAlcala, 2000). Draco spilopterus may warrant further
taxonomic attention once biochemical studies of species boundaries become available (Taylor, 1922c;
McGuire and Alcala, 2000).
Localities and specimens: . (Site 3) CAS 137578,
185504; (Site 4) PNM 1628–38; (Site 6) CAS
137608–609, 185505; (Site 7) PNM 1759–79; (Site 8)
CAS 127886, 127916, 127961, 128031; (Site 11)
PNM 1275–82, 2720–21; (Site 14) PNM 3769–70,
3878; (Site 18) CAS 127851–52; (Site 19) TNHC
58465–67, 58471–80, 58482–90, 58850; (Site 43)
CAS 39686; (Site 44) CAS 125295, USNM 38990–
96; (Site 47) CAS 125277–94.

Hydrosaurus pustulatus (Eschscholtz) (Figs.
29, 30)The Mt. Madja-as specimens were collected in overhanging stream-side vegetation. On Mt. Baloy we alsocollected a specimen in similar riparian habitats.
These large omnivorous agamids can be found on
trees and shrub-layer vegetation, overhanging streams
and rivers (Alcala, 1986). The taxonomy of Philippine
Hydrosaurus is in need further taxonomic studies.
Localities and specimens: (Site 11) CMNH 5043;(Site 13) TNHC 56762; (Site 22) USNM 77091–103,
85073–74; (Site 44) CAS 125336, USNM 38988–89;(Site 50) 77104–28, 78168–87.

Gonocephalus sp.
This genus was reported as an island record for Panay
from site 2 by Sison, et al. (1995). The name G.
sophiae has been applied to Negros populations (Taylor, 1922c; Alcala, 1986) and we might expect that if

Vol. 9, p. 13

Negros populations are indeed G. sophiae (and thisname applies to a distinct lineage that is independent
from G. semperi), specimens from Panay might bereferable to this species as well (see Gaulke, in press).
At present, no suitable taxonomic reference exists,and species boundaries are unclear. Due to this fact,
we do not apply a specific epithet to this population.
Philippine populations of the genus Gonocephalus are
greatly in need of taxonomic review.
Localities and specimens: (Site 2) PNM 1130–32;(Site 14) PNM 3858.

Gekkonidae
Cosymbotus platyurus (Schneider)
We found specimens of this common house gecko
species in both the city and in agricultural areas; they
are widely distributed on Panay (Brown and Alcala,1978) and are usually encountered under lights.
Localities and specimens: (Site 7) PNM 1803,
1815–16; (Site 8) CAS 128020; (Site 11) CMNH
5137, PNM 1261–74; (Site 19) CMNH 5089–96,
5098–104, PNM 2722–36; (Site 22) USNM 78776–
832, 103480; (Site 23) USNM 77144–57, 339998–
340010; CAS–SU 9613; (Site 26) PNM 1234–44,
1247, 1249; (Site 29) USNM 339995–97; (Site 34)
CAS–SU 9612, 12021; (Site 40) USNM 34011–12;(Site 50) FMNH 41302.

Cyrtodactylus annulatus (Taylor) (Fig. 32)
Brown and Alcala (1978) and Alcala (1986) report
this species in forested areas ranging from sea level to1200 m on the nearby land-bridge islands of Negros,Cebu, and Inampulugan (Site 42) and its presence on
Panay is not surprising. Nevertheless, this report andthat of Gaulke (in press) appear to be the first recordsof this species from Panay. Brown and Alcala (1978)
reported that this species has been collected on theforest floor, from under logs, beneath bark, and on thetrunks of trees. We found Panay specimens in secondary forest on root masses of trees protruding througheroding banks along a large river at the base of Mt.
Madja-as. Predation on this species by flying snakes(Chrysopelea paradisi) was observed in these same
microhabitats.
Locality and specimens: (Site 28) CAS 124614–15,
124768–78 (Site 42) CAS–SU 28009–10, 28013–14,
28016–19, 28031, 28036–41, 28044–46,28050–60.

Vol. 9, p. 14

Asiatic Herpetological Research

Cyrtodactylus philippinicus (Steindachner)(Fig. 33)
This species of Cyrtodactylus is found in a variety ofhabitats in the forest, particularly in rotting logs
(Brown and Alcala, 1978; Alcala, 1986) and has beencollected from sea level to nearly 1200 m (Brown andAlcala, 1978). In the Visayas, this species also has
been collected on the nearby land-bridge islands of
Negros, Pan de Azucar, and Boracay but never before
on Panay. The specimens from Mt. Madja-as werecollected in primary forest, during the day, underloose tree bark.
Localities and specimens: . (Site 6) CAS 137607;
USNM 496868; (Site 8) CAS 127883; (Site 11)
CMNH 5125–28, PNM 2751–53; (Site 13) TNHC
56339; (Site 18) 127864, 127869; (Site 28) CAS
124783–84; (Site 41) CAS 124046, 124780–82.

Gehyra mutilata (Weigmann) (Fig. 34)
This common and widely-distributed lizard is consistently found not only around human habitation (indarker areas, away from lights), but also in gardens
and forested areas on trees (Brown and Alcala, 1978;
Alcala, 1986).

2000(Site 28) CAS 124979; (Site 29) USNM 340016–17;(Site 31) PNM 1143; (Site 35) CAS 124393, 124318,
124866–75; (Site 37) CAS 124315–17, 124929–49;(Site 39) CAS 124389–92; (Site 41) CAS 124319,
124580; (Site 42) CAS-SU 27929; (Site 44) CAS
125251; (Site 46) CAS 125249–50; (Site 47) CAS
125247–48; (Site 49) CAS–SU 9589; (Site 50)
FMNH 41377–81, 41376.

Gekko gigante Brown and Alcala (Figs. 35,
36)
Gekko gigante was described by Brown and Alcala(1978) from the tiny land-bridge islands of GiganteNorth and Gigante South, off the northeast coast of
Panay (Fig. 2). This species has not been seen since
its 1968 discovery until a recent visit to the type locality by RMB and A. Alcala in June 2000. We found G.
gigante in small sea-side caves in karst limestone outcrops along the south coast of Gigante South. The
northeast coast is the best place to survey for this species on Panay if suitable limestone habitat can belocated.
Localities and specimens: (Site 35) CAS 124318,
124866–75 (Site 37) CAS 124315–17, 124929–49.

Localities and specimens: (Site 2) PNM 1682–86;(Site 3) CAS 137579; (Site 4) PNM 1649–53; (Site 7)
PNM 1824; (Site 8) CAS 127888, 127922–29,
127948–52, 127964, 127975–76, 127999–128000,
128054–55; (Site 11) CMNH 5105–16, 5198, PNM
1250–58, 2737–46; (Site 16) CAS 127504, 127607;(Site 18) CAS 127804, 127866; (Site 19) PNM 2737–
46; (Site 23) USNM 77158, 78834–35; (Site 26)
PNM 1245–49; (Site 28) CAS 124434–36, 124616–
19; (Site 29) USNM 340013–14; (Site 37) CAS
125029; (Site 39) CAS 124118–120, 124161–170;(Site 40) USNM 340015; (Site 41) CAS 124180–83;(Site 42) CAS-SU 28012, 28047–49; (Site 44) CAS
124505–510; (Site 46) CAS 124687–88; (Site 47)
CAS 124682–86, 125129–31; (Site 48) CAS 124511–
12; (Site 50) FMNH 41383.

Gekko mindorensis Taylor (Fig. 37)
Brown and Alcala (1978), and Alcala (1986) reportthat individuals of this species are common on wallsof caves, on tree trunks, and in leaves around the buttresses of trees. Elsewhere in the Visayas this species
has been reported from Negros, Cebu, and Caluya(Brown and Alcala, 1978). We have caught this species on cement walls, abandoned buildings in the forest, in road-cut culverts, and on large dead tree trunks
and stumps. Sison et al. (1995) reported this as anisland record from Site 29. The taxonomic distinctiveness of this species requires verification; recent datasuggest the widespread G. mindorensis may be conspecific with G. kikuchii from Taiwan (Crombie andOta, unpublished data; see also comment by Taylor,
1922c).

Gekko gecko (Linnaeus)
This common species is found around human habitation and in forest adjacent to disturbed areas. We collected specimens in secondary forest near the base of
Mt. Madja-as.

Localities and specimens: (Site 8) CAS 127882,
127884–85, 128021; (Site 11) One uncataloged specimen, deposited in PNM; (Site 16) CAS 127700; (Site18) CAS 127817; (Site 28) CAS 124767; (Site 29)
PNM 2500; (Site 39) CAS 124136.

Localities and specimens: (Site 1) PNM 1056–63;(Site 2) PNM 2667; (Site 7) PNM 1062–1063, 1792–96; (Site 8) CAS 128004; (Site 11) CMNH 5018–25,
PNM 1282, 1330, 2662–69; (Site 16) CAS 127582;(Site 17) CAS 127745; (Site 19) PNM 2665; (Site 23)
CAS–SU 9585–88; USNM 77142–43, 340018–19;

Hemidactylus frenatus Schlegel in Dumeril
and Bibron
This common house gecko is widespread on Panayand often is associated with Cosymbotus platyurus(Alcala, 1986) in well lighted areas in human habita-

2000

Asiatic Herpetological Researchtion. We collected several specimens near the base of
Mt. Madja-as on walls of houses in agricultural areas.
Localities and specimens: (Site 3) CAS 137580,
USNM 496869–70; (Site 7) PNM 1801–02, 1804–14;(Site 8) CAS 127878–81, 127897, 127913–14,
127930–45, 127953–58, 127963, 127977–78,
128001–03, 128022–26, 128034–35, 128056–57,
136742–44; (Site 11) CMNH 5148–54, PNM 1259–
60, 1646–48, 2771–76; (Site 16) CAS 127505–07,
127515–16, 127583, 127615–16, 127642–46,
127652–56, 127658–59, 127699; (Site 17) CAS
127710–20, 127727–44, 127748–65, 127795–99,
136741; (Site 18) CAS 127805–14, 127818–26,
127829–40, 127848–50, 127865; (Site 19) CMNH
5129–36, 5139–47, PNM 2754–70; (Site 23) USNM
78833, 78836–39; (Site 26) PNM 1245–1246; (Site28) CAS 124081–92, 124427, 124663, 125002–12;(Site 29) USNM 340021–27; (Site 33) USNM
340028–32; (Site 35) CAS 124363, 124504; (Site 37)124364–72, 124357–62, 124582–610, 124662; (Site39) CAS 124137–43, 124217–42; (Site 40) 340034–
41; (Site 41) CAS 124373–85, 124664–81, 125013;(Site 42) CAS–SU 28007–08, 28011, 28021, 28034–
35, 28042–43; (Site 44) CAS 124466, 124526–58[Nadulao Island]; (Site 46) CAS 124458–65, 125315–
30, 124736–66; (Site 47) CAS 124448–57, 125128,
125313–14; (Site 50) FMNH 41384–86.

Hemidactylus stejnegeri Ota and Hikida
Previously referred to H. garnoti (Brown and Alcala,1978), the status of Philippine populations was clarified by Ota and Hikida (1989) and Ota et al. (1993).
Sison et al. (1995) reported this species as an island
record for Panay.
Locality and specimen: (Site 25) CMNH 3225.

Hemiphyllodactylus insularis Taylor
While Brown and Alcala (1978) referred Philippine(and Panay Island) populations of Hemiphyllodactylus to H. typus, collections in USNM contain males
and are not, therefore, the widespread unisexual H.
typus, but probably are better referred to Taylor’s H.
insularis (R. Crombie, personal communication.).
Localities and specimens: . (Site 3) CAS 137581–
83; (Site 8) CAS 127889, 127965–71; (Site 18) CAS
127855–57.

Lepidodactylus lugubris (Dumeril andBibron)This lowland gecko is common in patchy distributions
and found in leaf axils, under bark, in tree holes, incoconut groves, and associated with large rocks in

Vol. 9, p. 15

coastal areas (Brown and Alcala, 1978; Alcala, 1986).
Though reported from Negros, Cebu, Inampulugan,
Boracay, Gigante, and Pan de Azucar (Brown andAlcala, 1978), this species has not yet been reported
from Panay.
Localities and specimens: (Site 8) CAS 127947,
127959, 127979, 127982–87, 127989, 128005–07,
128010, 128012–14, 128017, 128043–44, 128046–
50, 128059, 154716; (Site 28) CAS 124579; (Site 37)
CAS 124200, 125030, 125033; (Site 39) CAS
124116–17; (Site 41) CAS 124981–125000; (Site 42)
CAS–SU 28015, 28061; (Site 44) CAS 124561–63;(Site 46) CAS 124492.

Lepidodactylus planicaudus Stejneger
Alcala and Brown (1978) recorded this species incoconut trees, mangroves, and in fern axils; they havealso been taken on tree trunks in forests from sea levelto 700 m. Elsewhere in the Visayan Aggregate Island
Complex it is known from Cebu, Guimaras, Caluya,
Masbate and Boracay (Brown and Alcala, 1978).
Localities and specimens: (Site 3) CAS 139939;(Site 8) CAS 127921, 127980–81, 127988, 127990,
128008–09, 128011, 128015–16, 128018–19,
128045, 128051–53; (Site 16) CAS 127698; (Site 17)
CAS 127709; (Site 39) CAS 124115, 124134; (Site46) CAS 124823–26.

Scincidae
Brachymeles boulengeri taylori Brown (Figs.
38, 39)Usually encountered in agricultural areas (especiallycoconut plantations) adjacent to forest, this species
most often is collected from its preferred microhabitat, inside rotting coconut logs. It has also been collected in mature and secondary forest, from sea levelto 1200 m on numerous islands in the Visayas (Brown
and Alcala, 1980) but this is the first record fromPanay.
Localities and specimens: (Site 1) PNM 1148; (Site25) PNM 1148; (Site 39) CAS 124157; (Site 41) CAS
124044 (Site 42) CAS 27930–31, 27946–51, 27953,
27973–84, 27987–93.

Brachymeles talinis Brown
This skink occupies decaying vegetation and humus
on the floor of primary and, occasionally, secondary
forest (Brown and Alcala, 1980; Alcala, 1986). Alcala
and Brown (1980) reported its elevational range fromsea level to approximately 1200 m.

Vol. 9, p. 16

Asiatic Herpetological Research

Localities and specimens: (Site 6) 137603; (Site 8)
CAS 127870–75, 127962, 128029–30, 154689; (Site14) PNM 3852, 3856, 3859, 3909–10; (Site 16) CAS
127517–39, 127584–90, 127606, 127663–68; (Site28) 154692, 200522–31; (Site 39) CAS 124148,
200521; (Site 42) CAS–SU 27972, 27996–97; (Site50) CAS 137603.

Brachymeles tridactylus Brown (Fig. 40)
This species has been collected under logs, humus,
and other debris in forested regions from sea level toapproximately 900 m (Brown and Alcala, 1980; Alcala,1986). On Mt. Madja-as we found them under logs inwell-regenerated second growth forest below 200 m.

Localities and specimens: (Site 3) CAS 137566–75;(Site 6) CAS 137602, 137604; (Site 8) CAS 127876–
77, 127915, 127974, 128027–28, 128037–42; (Site11) PNM 5514–15; (Site 42) CAS–SU 27950, 27952,
27985–86, 28456.

Dasia grisea (Gray)In the Philippines, this species has been recorded fromthe islands of Mindoro, Luzon, and Marinduque(Brown and Alcala, 1980). Its inclusion in this paper
is based on a single record from Semirara Island, asmall island south of Mindoro that is included in thepolitical boundary of Antique province but is on theland-bridge to Mindoro. We do not expect this species
to be discovered on Panay and we do not regard it as a
Visayan Aggregate Island complex resident.
Locality and specimen: (Site 18) CAS 134218.

Dasia semicincta (Peters)
A large lowland species, previously reported only
from Mindanao in the Philippines (Brown and Alcala,1980; Alcala, 1986). Outside of the Philippines D.
semicincta is known from Borneo (Brown and Alcala,1980). This is the first record of this species fromPanay; it also is a substantial range extension in needof verification.
Locality and specimen: (Site 23) USNM 78840.

Emoia atrocostata (Lesson)
This species of skink is common in mangroves andcan be found active on tree trunks, in tree holes, and inrock crevices (Alcala and Brown, 1967; Brown andAlcala, 1980; Alcala, 1986). Taylor (1922c) reported
that this species swims in brackish water, burrows insand, and feeds on small crabs. On Panay and Negros,mangrove forests have nearly all been cleared, possibly rendering this species rare in recent collections.

2000

Localities and specimens: (Site 16) CAS 127638–
39; (Site 18) CAS 127842–45; (Site 22) CAS–SU
13585; (Site 23) USNM 77139, 78411, 78841, 80934;(Site 37) CAS 125040; (Site 41) CAS 124329–32,
124620–24; (Site 42) CAS-SU 27920; (Site 47) CAS
125335; (Site 50) USNM 77141.

Lamprolepis
smaragdina
philippinica(Mertens)
This skink is common in coconut groves and gardens
and a variety of natural vegetation types (Alcala,1986). Near Mt. Madja-as and Mt. Baloy we observed
them on coconut trunks at low elevations.
Localities and specimens: (Site 4) PNM 1639; (Site7) PNM 1780, 1782, 1786; (Site 8) CAS 127887,
127917–20, 127946, 127991–98, 128058, 128060–
61; (Site 11) CMNH 5038–40, PNM 1283, 2681–82;(Site 14) PNM 3798; (Site 16) CAS 127508, 127612–
14, 127640–41, 127669; (Site 17) CAS 127747,
127794; (Site 18) CAS 127853–54; (Site 19) TNHC
56472–73; (Site 28) 124053–57; (Site 35) CAS
124198, 124711–13; (Site 39) CAS 124201–02; (Site40) USNM 340061; (Site 42) CAS–SU 27954–62,
27969; (Site 44) CAS 125301; (Site 46) 125298–300;(Site 47) CAS 124447, 125296–97.

Lipinia pulchella taylori (Brown and Alcala)This widely distributed arboreal skink had not beenreported from Panay prior to our visits to Mt. Majdaas. Previously, L. p. taylori had only been known from
Negros (Brown and Alcala, 1980). Brown et al. (1996;
in press) have collected specimens from Luzon that
key out to this subspecies, potentially indicating theneed for a review of Brown and Alcala's (1980) taxonomy of this species. This is a new island record for
Panay.
Localities and specimens: (Site 11) CMNH 5083;(Site 25) PNM 1156.

Mabuya indeprensa Brown and Alcala
Sison et al. (1995) reported this species as an island
record, collected at about 200 m on Mt. Baloy. Well
known from forested, montane habitats on Negros andCebu (Brown and Alcala, 1980), this species is foundin leaf litter, stumps and fallen logs, and around tree
buttresses.
Localities and specimens: (Site 1) PNM 1149; (Site9) PNM 1153–55; (Site 11) CMNH 5119–22, 5199–
202, PNM 2747–49, 5511–13; (Site 13) one uncataloged specimen deposited in PNM; (Site 16) CAS
127596–97, 127599, 127601–02, 127604–05; (Site2000

Asiatic Herpetological ResearchVol. 9, p. 1718) CAS 127863; (Site 25) CMNH 3247, PNM 1151–
1152.

mens were collected in primary forest between 1400
and 1600 m in pit-fall traps.

Mabuya multicarinata borealis Brown andAlcala (Fig. 41)
This subspecies commonly is found under tree bark,logs and piles of vegetation (Brown and Alcala, 1980;
Alcala, 1986); in the Visayas it has been collected on
Negros, Caluya, Semirara, Gigante, Pan de Azucar,
and Cebu, but this is the first record from Panay.
Brown and Alcala (1980) reported finding one specimen as high as 1500 m on Luzon island.

Localities and specimens: . (Site 6) CAS 137644;(Site 11) CMNH 5062–74, PNM 2684–710; (Site 14)
PNM 3720, 3746–48, 3753, 3867–69, 3871, 3875,
3898, 3901–03; (Site 41) CAS 124048–49.

Localities and specimens: . (Site 2) PNM 1657–60,
1668–80; (Site 6) USNM 496871; (Site 16) CAS
127598, 127600, 127603, 127650–51; (Site 18) CAS
127862. 127867–68; (Site 28) CAS 124107–109,
124971–78; (Site 35) CAS 124493–97; (Site 36) CAS
124199, 125039; (Site 37) CAS 125038; (Site 41)
CAS 124050.

Mabuya multifasciata (Kuhl)
This species was found in a variety of habitats frombeaches and low elevation agricultural areas, to disturbed forest adjacent to primary forest on Mt. Madjaas. Though known to occur throughout the Philippines, in the Visayas, this species has only beenreported from Negros (Brown and Alcala, 1980); thisis the first published account from Panay.
Localities and specimens: (Site 2) PNM 1661–65;(Site 3) CAS 137585, USNM 496872; (Site 4) PNM
1640–45, 1655–56; (Site 6) CAS 137610–12, 139147;(Site 7) 1781, 1783; (Site 11) CMNH 5203–04, PNM
5510; (Site 14) PNM 3845–46, 3853; (Site 22) CAS–
SU 13632; (Site 25) PNM 1149–55; (Site 42) CAS–
SU 27968, 27971, 27998; (Site 44) CAS 125264–66;(Site 46) CAS 125334; (Site 47) CAS 125262–63,
125333; (Site 50) FMNH 41389–404.

Parvoscincus sisoni Ferner, Brown and Greer
All specimens of this recently-described species werecollected during the day from beneath loose soil andleaf litter in the forest between 900 and 1125 m(Ferner et al., 1997). At present, this species is known
only from Mt. Madja-as; the only other member of thegenus is from Palawan Island (P. palawanensis).
Locality and specimens: (Site 11) CMNH 3797–99,
PNM 2308–10.

Sphenomorphus arborens Taylor
This forest species is found under logs and leaves andoccasionally on buttresses at the base of tree trunks(Brown and Alcala, 1980; Alcala, 1986). Our speci-

Sphenomorphus coxi divergens Taylor
This species is often found at the forest edge and inforest gaps. It is widely-distributed in the archipelago(Brown and Alcala, 1980; Alcala, 1986). Sphenomorphus coxi divergens previously has been reported
from Luzon, Marinduque, and Mindoro but not the
Visayas (Brown and Alcala, 1980). This is the firstpublished record of a specimen from Panay.
Locality and specimens: (Site 11) CMNH 5123,
PNM 2750.

Sphenomorphus cumingi (Gray)
This large forest species is found around fallen logs
and tree buttresses and is believed to be limited to low
to medium elevations (Brown and Alcala, 1980;
Alcala, 1986). Sphenomorphus cumingi previously
has been reported from islands in the Luzon and Mindanao aggregate island complexes; this is the firstpublished record of a specimen from Panay.
Locality and specimen: (Site 28) CAS 124779.

Sphenomorphus jagori grandis Taylor (Fig.
42)
This large skink is found under rotting logs, in sun
spots in forested and disturbed areas, and along
streams (Brown and Alcala, 1980; Alcala, 1986).
Localities and specimens: . (Site 6) 137645–48;(Site 14) PNM 3724–5, 3755; (Site 16) CAS 127660–
62, 127677; (Site 28) CAS 125267–76, 154659–68;(Site 37) CAS 124835, 185508; (Site 41) CAS
185509–516.

Sphenomorphus steerei Stejneger
We found this species on Mt. Madja-as in leaf litter inmontane forest. Brown and Alcala (1980) report that
this species is common in leaf litter of primary forest
and in secondary growth.
Localities and specimens: (Site 3) CAS 137600–
601, 139169–70, USNM 496873–74; (Site 11)
CMNH 5026–37, PNM 2670–80; (Site 14) PNM
3708–09, 3721–23, 3726, 3749–52, 3754, 3775–81,
3786–87, 3797, 3826–29, 3836–42, 3847–51, 3854–
55, 3861, 3870, 3872–74, 3876–77, 3879–81, 3890–
92, 3914; (Site 25) PNM 1159; (Site 28) CAS

Vol. 9, p. 18

Asiatic Herpetological Research

124437–38, 124577–78; (Site 36) CAS 124830; (Site37) CAS 124827–29, 124839; (Site 39) CAS 125091;(Site 47) CAS 124625–61, 125064, 125090.

Tropidophorus grayi Günther (Fig. 43)
Sison et al.'s (1995) Mt. Baloy specimen was the firstrecord of this species from Panay. We also found a
specimen at the base of Mt. Madja-as underneath arock on the edge of a small stream in second growthforest.
Localities and specimens: (Site 11) CMNH 5117;(Site 13) One uncataloged specimen deposited in
PNM.

Varanidae
Varanus salvator nuchalis (Günther) (Figs. 44,
45)This monitor lizard is found in agricultural and disturbed areas, mangrove swamps, and forested areasfrom sea level to about 1200 m (Alcala, 1986; Gaulke,
1991a, 1991b, 1992). On Mt. Madja-as we collected
one specimen during the day in second growth forest
where it was active near a small stream.
Localities and specimens: (Site 1) PNM 1142; (Site11) PNM 5660; (Site 22) CAS 11018; (Site 31) PNM
1142; (Site 37) CAS 124879; (Site 44) CAS 124881;(Site 47) CAS 124880; (Site 50) USNM 77129–32,
80115–19; FMNH 41417–18.

Squamata (Snakes)
Acrochordidae
Acrochordus granulatus (Schneider)
Found in mangroves and at river mouths, this species
feeds exclusively on fish and is widely distributed andcommon (Alcala, 1986). We are not aware of otherpublished accounts of this species from Panay.
Locality and specimens: (Site 23) USNM 78412,
78744–45, 78906, CAS–SU 8695, 8769.

BoidaePython reticulatus (Schneider) (Fig. 46)
While this snake is considered widely distributed andcommon in the tropical rain forests as well as nearhuman habitation (Alcala, 1986), only one specimen
has been collected on Panay Island (Leviton, 1963c;
see also Gaulke, in press).
Locality and specimen: (Site 44) CAS 124916.

2000

Colubridae
Ahaetulla prasina preocularis (Taylor) (Fig.
47)
This snake may be found in low trees and shrubs fromsea level to about 800 m on Panay (Leviton, 1963c,
1968; Alcala, 1986). The specimens from Mt. Madjaas were collected from vegetation overhanging asmall stream at low elevation.
Localities and specimens: (Site 2) PNM 1689; (Site7) PNM 1787; (Site 11) CMNH 5084–85, PNM 2716;(Site 14) PNM 3825; (Site 29) USNM 340042; (Site44) CAS 125339; (Site 50) FMNH 41108.

Boiga angulata (Peters)
We found our specimen in a coconut tree in a clearing
adjacent to virgin forest at 900 m on Mt. Madja-as.
While this species is known from Negros (Leviton,
1970a), until this report and that of Gaulke (in press)
it has never been recorded on Panay.
Locality and specimen: (Site 11) CMNH 5504

Boiga cf cynodon (Cuvier in F. Boie)
This species has been found in forested areas at low
altitudes on Palawan, Mindanao, and Luzon (Leviton,
1963c, 1970a; Alcala, 1986) but previously it has notbeen reported in the Visayas (see also Gaulke, inpress).
Locality and specimen: (Site 41) CAS 125173.

Boiga cf. dendrophila (Boie)
Mangrove snakes usually are found in branches of
low trees and bushes in forested areas (Leviton 1968;
Alcala, 1986) at low elevations. This is the first recordof a species in this complex from Panay.
Localities and specimens: (Site 28) CAS 124388;(Site 39) CAS 124386–87.

Calamaria geravaisi Dumeril and Bibron
A burrowing snake, this species is commonly found in
the humus under rotting logs and feeds on earthworms(Leviton, 1963c; Inger and Marx, 1965; Alcala, 1986).
On Mt. Madja-as we found specimens under rotten
logs and large flat rocks.
Localities and specimens: (Site 2) PNM 1087; (Site11) CMNH 5081–82, PNM 2714–15; (Site 23) CAS–
SU 15953–57, 15962–65; (Site 44) CAS 124612.

2000

Asiatic Herpetological ResearchVol. 9, p. 19

Cerberus rynchops (Schneider)This aquatic snake has been collected in brackish
swamps, mangroves, fish ponds, and river estuaries incoastal areas (Gyi, 1970; Alcala, 1986)

127707–708; (Site 18) CAS 127828, 127847, 127859;(Site 28) CAS 125170; (Site 39) CAS 124203,
185673–74; (Site 42) CAS-SU 28004; (Site 50)
FMNH 41093–96.

Localities and specimens: (Site 1) PNM 1053–55,
1077–83; (Site 20) CAS–SU 13079; (Site 22) CAS–
SU 8696–97, 8719; CAS–SU 12380, CM R2423,
R2426; (Site 23) USNM 77159–478, 78907–19; (Site40) 340043; (Site 50) FMNH 41115–17.

Dendrelaphis pictus pictus (Gmelin) (Fig. 48)
This common snake is arboreal and usually foundnear streams, in vegetation surrounding flooded rice
fields, and in swampy areas (Leviton, 1963c, 1970b;
Alcala, 1986). The Mt. Madja-as specimens were collected at night where they slept in stream-side vegetation.

Chrysopelea paradisi Boie
This species was not known from Panay (Leviton,
1963, 1964a) until Sison, et al. (1995) reported it as
an island record from Site 1. On Mt. Madja-as wefound this arboreal snake during the day in root
masses overhanging a river bank; one specimen waspreying on a Cyrtodactylus annulatus when captured.
We have also observed this species in coconut grovesat sea level near the town of San José (site 19).
Localities and specimens: (Site 1) PNM 1050; (Site8) CAS 128032; (Site 11) CMNH 5041–5042, PNM
2683; (Site 19 TMM 56474; (Site 27) CAS 185–672;(Site 36) CAS 125172; (Site 46) CAS 125331–32.

Cyclocorus lineatus alcalai Leviton
We found our specimens in habitats ranging from disturbed second growth at sea level to first growth forested riparian sites at 1400 m. At the time of its
description (Leviton, 1967), this subspecies was
known only from the nearby islands of Negros andCebu (Fig. 2). This is the first published account of
this species from Panay.
Localities and specimens: (Site 1) PNM 1047,
1065; (Site 2) PNM 1065, 1688; (Site 3) CAS
137576; (Site 6) CAS 137606; (Site 11) CMNH
5086–87, PNM 2717–18; (Site 14) 3884–85, 3912;(Site 16) CAS 127702, 127706; (Site 28) CAS
124051–52, 124421; (Site 37) CAS 125171; (Site 47)
CAS 124445.

Dendrelaphis
caudolineatusterrificus(Peters)
This subspecies is known from Panay and Negros andusually is found in forested and cultivated areas in oralong swamps and streams (Leviton, 1970b; Alcala,1986). Sison et al. (1995) reported this as an island
record from 200 m on Mt. Baloy. The specimen from
Mt. Madja-as was sleeping in bushes near a riverwhen captured.
Localities and specimens: (Site 4) CAS 1654; (Site8) CAS 127896, 128033; (Site 11) CMNH 5080; (Site13) CMNH 3254; (Site 16) CAS 127703–705,

Localities and specimens: (Site 1) PNM 1049,
1071; (Site 3) CAS 137577; (Site 7) PNM 1788–90;(Site 9) PNM 1071; (Site 11) CMNH 5078–79, PNM
2713; (Site 14) PNM 3843–44; (Site 22) CAS–SU
8660–70, 8718, 14932, 14936–37, CMNH 2408–13,
CM R2226, R2408–13; (Site 23) CAS–SU 8698–99,
8708–710; USNM 77419–591, 340044–51; (Site 30)
CAS–SU 14931 ; (Site 38) CAS 200256; (Site 44)
CAS 125255–61; (Site 46) CAS 125254; (Site 47)
CAS 125252–53; (Site 48) CAS 124725; (Site 50)
FMNH 4109–104, 41106, USNM 77592–609.

Elaphe erythrura psephenoura Leviton
This is a common lowland snake, often found nearhuman habitations (Leviton, 1979; Alcala, 1986). On
Mt. Madja-as we collected one specimen from the forest floor where it was active at midday.
Localities and specimens: (Site 1) PNM 1048,
1051; (Site 11) PNM 5662; (Site 22) CAS–SU 12389,
13212–13; (Site 23) USNM 340052; (Site 30) CAS–
SU 13217; (Site 38) CAS 131700; (Site 42) CAS-SU
28001; (Site 47) CAS 125141–42, 125340.

Gonyosoma oxycephala (Reinwardt in F.
Boie)This is an arboreal snake that is found in disturbed
and primary forests (Alcala, 1986). On Mt. Madja-as
we collected one specimen that was active at mid day
in a tree above a large river (4 m from the ground).
This record, and that of Gaulke (in press) are the firstpublished accounts of this species from Panay.
Localities and specimens: (Site 1) PNM 1084; (Site9) PNM 1084; (Site 11) CMNH 5503; PNM 5659.

Hologerrhum dermali Brown, Leviton, Ferner,and Sison
We first collected this newly-described species(Brown et al., this issue) between 1030 and 1510 m inclimax forest on Mt. Madja-as. Specimens were collected in a dry stream bed and in leaf litter on the for-

Vol. 9, p. 20

Asiatic Herpetological Research

est floor 30 m from a large river. One specimen from
the Municipality of San Remegio was collected at
approximately 700 m above sea level. Recent survey
work in NW Panay suggests that this species also
occurs in forested areas at lower elevations (Gaulke,in press).
Localities and specimens: (Site 11) CMNH 5075,
PNM 2711 (14) PNM 3704.

Lycodon aulicus capucinus (H. Boie in F.
Boie) (Fig. 49)
This common, widespread, nocturnal snake is often
found in gardens, agricultural areas, and aroundhouses (Leviton, 1965; Alcala, 1986). Our specimens
were collected near rice fields away from forested
areas.
Localities and specimens: (Site 1) PNM 1072–75;(Site 3) CAS 137584; (Site 8) CAS 127960; (Site 12)
PNM 1072; (Site 13) PNM 1380; (Site 22) CAS–SU
8671, 8700–01, CMNH 2443, CM R2443; (Site 23)
USNM 77616, 340053; (Site 26) PNM 1073–75; (Site47) CAS 125341.

Oligodon modestum (Günther)
This species is found under rotting logs and forest
floor debris; it is known from sea level to 400 m(Alcala, 1986). In the Visayas, this species was only
known from Negros (Leviton, 1963a) before Sison etal. (1995) first reported it on Panay.
Localities and specimens: (Site 1) PNM 1066; (Site2) PNM 1067; (Site 14) PNM 3790, 3866.

Psammodynastes pulverulentus (H. Boie in F.
Boie)
Leviton (1963c; 1983) and Alcala (1986) report thissnake as common up to elevations of 1000 m or morein moist forests on Negros Island; Leviton’s (1963c)
listing of this species from Panay was not repeated in
his 1983 review of the genus in the Philippines and no
specimens were reported from Panay in the later
paper (Leviton, 1983). This account and that of
Gaulke (in press) appears to be the first vouchered
records of this species from Panay.
Localities and specimens: (Site 1) PNM 1068; (Site2) PNM 1069; (Site 25) PNM 1067, 1070.

Pseudorabdion mcnamarae Taylor
Prior to Sison et al. (1995) this species was known
only from Negros and Luzon (Brown and Leviton,
1959; Alcala, 1986). Specimens from Mt. Baloy werecollected at 950 m under rotting logs in original forest.

2000

Locality and specimens: (Site 13) Two uncataloged
specimens in PNM (PNM Field Numbers 163 and209).

Pseudorabdion oxycephalum (Gunther)Previously considered a rare snake endemic to Negros
Island, this species is now known from other localities
in the Luzon and Mindanao aggregate island complexes (Brown and Leviton, 1959; Leviton, 1963c;
Alcala, 1986; Brown et al., 1999). This is the firstpublished record from Panay.
Pseudorabdion
oxycephalum is been found in humus and under rotting logs from sea level to about 750 m.
Localities and specimens: (Site 6) CAS 137643;(Site 39) CAS 124174, 124193; (Site 41) CAS
124043.

Pseudorabdion talonuran Brown, Leviton andSison
The discovery of this new species at high elevations
was surprising. Both specimens were found underlogs on Mt. Madja-as in forest classified as the transition zone between mixed dipterocarp (submontane)
and mossy (upper montane; Whitmore, 1984). The
holotype was found at 1500 m and the paratype at1410 m.
Locality and specimens: (Site 11) CMNH 5076,
PNM 2712.

Tropidonophis negrosensis (Taylor)
This species of water snake is common along forest
streams from sea level to about 700 m (Leviton,
1963c; Alcala, 1986; Malnate and Underwood, 1988).
The specimen from Mt. Madja-as was found on ariver bank at midday.
Localities and specimens: . (Site 3) CAS 185749;(Site 6) CAS 137613; (Site 11) CMNH 5124; (Site14) PNM 3911; (Site 22) CAS–SU 15971; (Site 28)
CAS 124611; (Site 41) CAS 124047.

Zaocys luzonensis Gunther
A common tropical forest snake, this species ranges
from sea level to over 1100 m (Leviton, 1983; Alcala,1986; Ross et al., 1987). On Mt. Madja-as, we found
two specimens active at midday in disturbed forest at
low elevation.
Localities and specimens: (Site 1) PNM 1052; (Site2) USNM 269078; (Site 7) PNM 1791; (Site 11)
CMNH 5505; PNM 5663.

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ElapidaeCalliophis calligaster gemianulis (Peters)
We collected one specimen on Mt. Madja-as at
approximately 800 m on a mountain path away fromwater. They have previously have been found in avariety of semifossorial habitats associated withregenerated and climax forest (Leviton 1963b, 1963c;
Alcala, 1986)
Localities and specimens: (Site 11) CMNH 5088,
PNM 2719; (Site 13) PNM 1379; (Site 21) UPLB
2184; (Site 22) CM R2581; (Site 23) CAS-SU 12966–
68; (Site 44) CAS 125364; (Site 47) CAS 125363.

Hydrophis belcheri Gray
This species has been recorded from the Visayan sea
and is thought to be rare (Alcala, 1986).
Localities and specimens: (Site 5) USNM 38588;(Site 37) FMNH 202832–36, 202838, 202840–42.
Hydrophis cyanocinctus Daudin

Alcala (1986) mentioned records from the
Visayan Sea.
Locality and specimens: (Site 37) FMNH
202843–54.
Hydrophis elegans Gray
Locality and specimen: (Site 5) USNM 38589.

Hydrophis inornatus Gray
Locality and specimen: (Site 23) CAS-SU 8778.

Lapemis hardwickii Gray
Alcala (1986) reported this species from the Visayansea; he noted that specimens were collected by trawling.
Localities and specimens: (Site 23) USNM 77610–
15.

Laticauda colubrina Schneider
This sea snake is found among rocks and in coral reefareas near shore; it is commercially used for its meatand hide (Alcala, 1986).
Locality and specimens: (Site 37) FMNH 202797–
801, 202804–808, 202810.

Typhlopidae
Ramphotyphlops braminus (Daudin)This species is common under rocks and other debris
in agricultural areas. It can also be found in similar

Vol. 9, p. 21

microhabitats along the edges of forested areas(Alcala, 1986). Taylor (1922d) reported catching
many specimens under rocks immediately following
rains.
Localities and specimens: . (Site 1) PNM 1044–46;(Site 8) CAS 127972, 128036; (Site 17) CAS 127746;(Site 18) CAS 127846, 127858, 127860; (Site 30)
CAS–SU 12544–49; (Site 37) CAS 125041; (Site 46)
CAS 124503.

Ramphotyphlops cumingii (Gray)
Alcala (1986) reports this blind snake is found in epiphytic ferns in the trees of moist forests at low to
medium elevations. In all of his field work, Taylor(1922d) was unable to capture this species, suggesting
he may have overlooked its preferred microhabitat.
Localities and specimens: (Site 28) CAS 169877;(Site 35) CAS 125092.

Typhlops castanotus Wynn and Leviton
Described very recently (Wynn and Leviton, 1993),
this distinctive bicolored species is known from
Negros, Panay (Makato), Boracay, and Inampulugan
islands. It has been collected under debris in bamboo
and coconut groves, hardwood forests and forest remnants.
Localities and specimens: (Site 3) CAS 139171;(Site 8) CAS 127973; (Site 42) CAS–SU 27934–46.

Typhlops hypogius Savage,Typhlops luzonensis Taylor, andTyphlops ruber Boettger
Found in detritus under rotting logs, these snakes havebeen collected in forests and disturbed areas near forest from sea level to about 800 m (Alcala, 1986).
Wynn and Leviton (1993) followed McDowell (1974)
in referring Philippine T. luzonensis and T. hypogius
to T. ruber. Recently, McDiarmid et al. (1999)
asserted that T. luzonensis and T. hypogius should berecognized until more compelling evidence is presented that would suggest they are conspecific with T.
ruber. We agree, noting that the type localities for T.
hypogius (Cebu), T. ruber (Samar), and T. luzonensis(Luzon) are each located on separate Pleistocene
aggregate island complexes that might be expected tosupport independent lineages of blind snakes. If this isso, and the Visayan islands contain a single distinct
lineage in this species complex, the name Typhlops
hypogius would most likely apply to specimens from
Cebu, Negros, Panay, Masbate, and smaller, landbridge islands.
Localities and specimens: (Site 18) CAS 127861.

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Family Viperidae
Tropidolaemus cf wagleri (H. Boie in F. Boie)(Figs. 51, 52)
This arboreal viper is common in forest bushes andsmall trees as well as mangroves (Taylor, 1922d; Leviton, 1964b; Alcala, 1986). The Mt. Madja-as specimen was found in a banana plantation at 800 m near
primary forest. We find it unlikely that all SE Asian
populations currently referred to T. wagleri will prove
to be a single species. This is the first account of a
specimen referable to this species from Panay.
Localities and specimens: (Site 11) CMNH 5076.

Discussion
The primary goal of this report has been to provide acomprehensive and synthetic review of the amphibianand reptiles species known from Panay Island and issurrounding land-bridge islets. As noted, over the past
century, biologists have expected that the documented
presence of a species on the neighboring island of
Negros implied its undocumented presence on Panay
as well (see Inger, 1954; Brown and Alcala, 1970,
1978, 1980, 1986; Brown and Rabor, 1967; Alcala,1986). However, Panay (and its smaller satellite islets)
supports low levels of endemicity, primarily as exemplified by populations at higher montane elevations.
Panay has several species of amphibians and reptiles
that have not yet been reported on Negros, whichalong with Cebu, is known to support species that so
far have not been documented on Panay. The presence
on Panay of numerous new and undescribed species
suggests that calculations of Panay’s herpetologicalendemicity will continue to rise with continued survey
work. Additionally, basic taxonomic studies of species complexes with representatives on Panay will no
doubt further contribute to the total number of Panay
endemics.
There are only a few reports of amphibians andreptiles of high elevation forests of Panay (Sison etal., 1995; W. Brown et al., 1997a; Ferner et al., 1997;
R. Brown et al. 1999, this issue). Essentially, on eachoccasion that herpetologists have surveyed higher elevation forests of Panay, they have discovered new species. Other surveys conducted in forested regions ofPanay include survey efforts of A. Diesmos, R. Crombie, and M.Gaulke (in press). Further high elevation
surveys in well forested regions of Panay are greatly
needed to gain an understanding and appreciation ofthese presumably relictal faunal elements.
There are numerous records that are included inthis report that were not at all unexpected. These

2000

include widespread SE Asian and Philippine species
that biologists have expected or assumed were present
on Panay. Others have even been listed as known fromPanay, but without specific reference to reliable locality data or museum specimens (see Alcala, 1986).
These include the frogs Bufo marinus, Kaloula picta,
Occidozyga laevis, Platymantis corrugatus, Platymantis dorsalis, Rana vittigera, the skink Mabuyamultifasciata, and the snakes Ramphotyphlops braminus, Acrochordus granulatus, Psammodynastes pulverulentus,
Gonyosoma
oxycephala,andTropidolaemus cf wagleri.
Another group of species includes forms known
from other well-surveyed islands in the Visayas(Negros, Cebu) that we expected to (and did) find on
Panay as well. Their presence on Panay was almost acertainty based on known biogeographic relationshipsof these islands. These species include the frogs
Kaloula c. negrosensis, Limnonectes cf leytensis, theskinks Lipinia pulchella taylori, Brachymeles boulengeri taylori, Mabuya multicarinata borealis, Tropidophorus grayi, Emoia atrocostata, the geckos
Cyrtodactylus annulatus, Cyrtodactylus philippinicus,
the sail-fin agamid Hydrosaurus pustulatus, and thesnakes Boiga angulata, Boiga cf cynodon, Boiga cf
dendophilia, Cyclocorus lineatus alcalai, Pseudorabdion mcnamarae, Oligodon modestum, and Trimereserus flavomaculatus (Gaulke, in press).
Several other records are major range extensions
and real surprises. The skink Dasia semisincta is otherwise known only from Borneo and Mindanao Islandand its presence on Panay is based on a wellvouchered specimen (USNM 78840) that can not be
discounted. The presence of a new species in thegenus Hologerrhum was surprising in that this genus
previously was considered a monotypic Luzon Aggregate Island Complex endemic (Leviton, 1963c; Brown
et al., this issue; Gaulke, in press). A forest species of
Kaloula related to K. kalingensis and K. kokacii is thefirst record of this species group outside the boundaries of the Luzon Aggregate Island Complex (Inger,
1954; Brown and Alcala, 1970; Alcala and Brown,
1998; Brown and Diesmos, unpublished data; Gaulke,in press). Some records (based on few specimens or
dubious locality data) may be in error; these include
Sphenomorphus coxi divergens, Sphenomorphus
cumingi, and S. fasciatus (of Sison et al., 1995); otherwise, if accurate, these records represent major
range extensions beyond the confines of the Mindanao, Mindoro, and Luzon aggregate island platforms.
As mentioned, a number of new species recently
have been described as Panay endemics; the presenceof endemics in high elevation habitats of Panay was

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not unexpected. These species Hologerrhum dermali(Brown et al., this issue; see also Gaulke, in press),
Parvoscinus sisoni (Ferner et al., 1997), Platymantis
panayensis (Brown et al., 1997a), Pseudorabdion talonuran (Brown et al., 1999), Kaloula sp., Platymantis
sp. 1, Platymantis sp. 2, and Platymantis sp. 3 (thisreport, see also Gaulke, in press).
Finally, there are numerous “subspecies”, species,
or members of widespread Visayan, Philippine, or SE
Asian species complex members on Panay that are of
uncertain taxonomic status and in need of basic taxonomic review (see individual species accounts). Many
species currently listed from Panay are, we expect,distinct lineages that eventually will be recognized as
Panay or Visayan endemics. These species are too
numerous to list and extend from common, low elevation forms to rare high elevation forest obligates.
There is much basic taxonomic work still to be conducted on Panay, in the Visayas, and in the rest of thePhilippines and we expect our estimates of Panay’s
endemicity to generally rise with further systematicstudies of the herpetofauna of the Philippines.
Faunal similarity calculations (Fig. 53) indicatedthat, as expected, the Visayan islands of Negros andCebu (situated on the same Pleistocene aggregate
island platform as Panay) were among the islands faunistically most similar to Panay (Masbate was notconsidered due to the absence of substantial records
from this island). We interpret this as evidence formid- to late-Pleistocene land bridge connections
between these islands (see Heaney, 1985, 1986).
Within the Visayan Aggregate Island complex, estimates of amphibian faunal similarities exceeded thoseof reptiles but when Panay is compared to islands outside the Visayan Aggregate Island complex, reptile
faunal similarities exceed those of amphibians. As
expected, amphibian faunal similarity between Luzon
and Panay is much lower than estimates calculated for
Cebu or Negros and Panay. However, surprisingly,
Luzon and Panay had a higher reptile coefficient ofsimilarity than did Cebu and Panay. A portion of thisunusual finding may be the result of sampling error
reflecting the degree to which survey data for Negros,
Panay, and Luzon are available. Calculations of similarities between Negros and Luzon (not shown) arehigher in reptiles (0.58) than they are for amphibians(0.45).
In comparisons with islands on other (nonVisayan) major Philippine aggregate island platforms,
reptilian species similarity was consistently higher
than that of amphibians. This may be due to antiquated taxonomy. Recently, disproportionate amountsof taxonomic work has been conducted on amphibianVol. 9, p. 31

groups, resulting in the recognition of more amphibian endemics, with fewer "shared" species among
islands. In contrast, many reptiles "species" are shared
between Panay and Luzon. Some of these may represent species complexes in need of taxonomic resolution. Alternatively, these calculations may reflect therelatively greater dispersal abilities presumed for reptiles when crossing ocean barriers. It is tempting toconsider that more reptile species may be shared
between these islands because reptiles may be moretolerant of exposure to heat and salt water, and may
have a higher probability of surviving dispersal events(e.g., via rafting) than would be expected for amphibians. In general, these results support the suggestionthat the herpetofaunal communities of islands within
the Visayan Aggregate Island Complex are very similar but also that they have their own degree of endemism and are far from being identical.
Species that we might expect to soon be discovered on Panay include populations that are otherwise
known from Negros and Cebu islands. However, wenote that the high elevation endemics of Negros (e.g.,
Platymantis hazelae, Pseudorabdion montanum) arenot expected to be discovered on Panay; in their place
we expect Panay to support it’s own high elevation
populations of closely-related montane endemics(e.g., Platymantis panayensis, Pseudorabdion talonuran). These include the frogs Rhacophorus pardalis(Fig. 54), Rhacophorus appendiculatus (widespreadon Mindanao and Luzon aggregate island complexes;
both known from Negros), Platymantis spelaeus(known from Negros; Fig. 55), geckos such as Lepidodactylus lugubris (widespread in the Philippines),
Lepidodactylus herrei (currently comprised of two
subspecies: L. h. herrei on Negros and L. h. medianus
on Cebu), Lepidodactylus christiani (known from
Negros and Cebu), Pseudogekko brevipes (known
from Negros and Cebu; P. compressicorpus has beencollected on Masbate), a Luperosaurus species (L.
cumingi has been collected on Negros), and skinks
like Lipinia quadrivittata quadrivittata (from Negros
and Cebu), Lipinia auriculata auriculata (fromNegros and Masbate), and Lipinia rabori (fromNegros). Snake species we expect will be found on
Panay with continued survey efforts include forms
possibly related to T. canlaonensis, T. hedraeus(Negros forms), Oxyrhabdion leporinum visayanum(from Negros and Cebu; Fig. 56), and Ophiophagus
hannah (recorded from numerous islands in the Philippines). The Philippine endemic crocodile, Crocodylus mindorensis, may have recently been rediscoveredon Negros (E. Alcala, personal communication) andmay be present on Panay if suitable habitat can beVol. 9, p. 32

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Figure 53. Faunal similarity coefficients, calculated between Panay and other majorislands in the Philippines. See text for formula and discussion.

located. As noted, the Gigante Island endemics Platymantis insulatus and Gekko gigante might be
expected to occur on karst limestone outcrops on
Panay’s northeast coast.
The future of exploration on Panay guarantees
continuation of the kind of discovery reported here.
There is an immediate need for continued basic survey efforts in the montane portions of Antique, Aklan,
and Iloilo provinces, all of which contribute to thewestern coastal mountain range that supports so muchof Panay’s herpetological endemicity. Additionally,
low elevation portions of Panay (principally Capizand Iloilo provinces) are also herpetologicallyunknown. If areas of overlooked primary forest or
well-regenerated secondary forest can be located, we
have high expectations that these will support novel
herpetological communities and generate continued
discoveries of new taxa. Areas of particular interest
include karst limestone outcrops along the northernand eastern coasts, mangrove fragments, cave habitats, and isolated outcrops of moderate elevation ineastern Panay (Fig. 2). As noted, the best place tosearch for the Gigante endemics Platymantis insulatus and Gekko gigante is limestone outcrops supporting caves along Panay’s northeastern coast. Other rare
Negros species (i.e., Luperosaurus cumingi, Lepidodactylus herrei, Platymantis spelaeus) may eventually
be revealed on Panay as well, once adequate surveys
in preferred microhabitats (forest canopies and karst
limestone caves; C. N. Dolino, personal communication) become available.

Like most islands in the Visayan Aggregate Island
Complex, Panay should be regarded a priority for
future conservation initiatives and programs aimed atsustainable resource management. Panay is a uniqueisland (not at all identical to Negros) that deserves its
own conservation efforts.

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The few remaining forests of Panay continue to befelled at an alarming rate, suggesting that its endemic
flora and fauna may disappear before even being
recorded by biologists. Low elevation forests andmangroves are all but gone, and even the most disturbed and negatively impacted sites warrant immediate study of the kind that currently is underway on
Cebu and Negros (A. Alcala, and E. Alcala, C. N.
Dolino, J. C. Gonzales, and M. Pedregosa, personalcommunication). It is our hope that Panay will be recognized as a model island ecosystem, ripe for collaborative efforts of conservation biologists, taxonomists,biogeographers, community organizers, and politicians. Conservation efforts targeted at the community
level represent the best opportunity for foreign, government, and non-government organizations in theireffort to halt the destructive practices of non-sustainable timber and mineral extraction industries that currently operate unchecked in the central Visayan
islands of the Philippines.

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