a new species of limestone-forest frog, genus...

A NEW SPECIES OF LIMESTONE-FOREST FROG, GENUSPLATYMANTIS (AMPHIBIA: ANURA: CERATOBATRACHIDAE) FROM

EASTERN SAMAR ISLAND, PHILIPPINES

CAMERON D. SILER1,3,4, ANGEL C. ALCALA

3, ARVIN C. DIESMOS2,3, AND RAFE M. BROWN

1,3

1Natural History Museum and Biodiversity Research Center, Department of Ecology and Evolutionary Biology,University of Kansas, Lawrence, KS 66045-7561, USA

2Herpetology Section, Zoology Division, Philippine National Museum, Rizal Park, Burgos St., Manila, Philippines3Angelo King Center for Research and Environmental Management, Silliman University, Dumaguete City, 6200,

Philippines

ABSTRACT: A new species of forest frog of the genus Platymantis is described from an elevation of 140 min the Taft Forest Reserve in eastern Samar Island, Philippines. It is assigned to the Platymantis guentheriSpecies Group, a group of primarily arboreal species, and is distinguished from these and other congeners byfeatures of its external morphology and its preferred terrestrial, limestone microhabitat. Several strikingmorphological characters include a large body (34.2–39.1 mm SVL for 9 males and 44.3–49.8 mm SVL for 9females), greatly expanded finger and toe discs, large eyes, spotted flanks, and sparsely-distributed, salmon-colored dorsal dermal tubercles. The new species represents the second largest Philippine Platymantis, thethird terrestrial species in the P. guentheri Group, and the only known species of Platymantis from theMindanao Faunal Region with a preference for forested, karst habitats.

Key words: Biodiversity; Cryptic species; Endemism; Faunal Region; Limestone frogs; Platymantisguentheri Species Group; SW Pacific

FROGS of the genus Platymantis have twogeographic centers of diversity and endemism.One large group of species is situated in theSolomon Islands (W. C. Brown, 1952, 1997;Menzies, 2007) and nearby Bismarck (Zweifel,1960, 1975; Foufopoulos and R. M. Brown,2004; R. M. Brown et al., 2006a,b; Foufopou-los and Richards, 2007) and Admiralty archi-pelagos (26 species; Allison, 1996; Richards etal., 2007). Another large group of speciesoccurs in the Philippines (27 species; Alcalaand W. C. Brown, 1998, 1999; W. C. Brown etal., 1997a; R. M. Brown and Gonzales, 2007).Outside of these two large archipelagos, onespecies occurs in Palau (Crombie and Pregill,1999); seven or eight are known from NewGuinea faunal region (Zweifel, 1969; Allison,1996; Gunther, 1999, 2005; Menzies, 2007);two to four species occur in eastern Indonesia(Edgar and Lilley, 1993; Menzies, 1982a,b;including undescribed taxa; M. I. Setiadi andD. T. Iskandar, personal communication); andtwo others occur in Fiji (Gorham, 1965;Morrison, 2003).

Within the Philippines, three speciesgroups are recognized on the basis of external

morphology (W. C. Brown et al., 1997a,c;Alcala and W. C. Brown, 1999)—the P.dorsalis Group, the P. hazelae Group, andthe P. guentheri Group. In addition to themorphological variation among Platymantis inthe Philippines, the advertisement calls andpreferred microhabitats of the species alsovary (R. M. Brown et al., 2002; R. M. Brownand Gonzales, 2007). Surveys have shown thatlimestone-forest habitats of the Philippinespossess highly distinctive species of Platyman-tis (W. C. Brown and Alcala, 1982; R. M.Brown and Alcala, 2000; Siler et al., 2007).

In 1998 the late Walter C. Brown (seeAlcala, 2004) informed RMB about a pair ofhighly distinctive Platymantis specimens fromthe Island of Samar that he had examinedmany years prior. He remarked that thesuspected new species was ‘‘a giant form thatlooked like a cross between Platymantisspeleaus and Platymantis insulatus.’’ AlthoughBrown could not remember where the spec-imens were deposited, his verbal descriptionof the suspected new species of giant Platy-mantis from Samar was quite specific and verydetailed. Over the past decade we have beenunable to locate the specimens in U.S.collections, but we have anxiously awaitedthe discovery of the new species that Brown4 CORRESPONDENCE: e-mail, [email protected]

Herpetologica, 65(1), 2009, 92–104

E 2009 by The Herpetologists’ League, Inc.

92

predicted, ‘‘would eventually turn up onSamar.’’

During the month of October 2007, weconducted herpetological field surveys in thelower-elevation forest of the Municipality ofTaft in eastern Samar Island (Fig. 1). Twenty-one individuals of an undescribed species ofPlatymantis that match Walter Brown’s de-scription were collected at an elevation of140 m. Despite its striking tree frog appear-ance, the new species is found in terrestrialmicrohabitats in karst forest, an ecomorpho-logical combination only shared with P.insulatus (R. M. Brown and Alcala, 2000).Herein, we describe the new species andreport on its natural history, ecology, andhabitat.

MATERIALS AND METHODS

We recorded morphometric data fromalcohol-preserved specimens that were fixedin 10% formalin (Appendix 1). Sex wasdetermined by gonadal inspection, and mea-surements were taken with digital calipers tothe nearest 0.1 mm. To minimize inter-

observer bias and other sources of potentialerror (Hayek et al., 2001; Lee, 1982), allmeasurements were scored by CDS. Descrip-tions of coloration were based on digitalphotographs and field notes of CDS.

Characters measured follow the definitionsof Siler et al. (2007) and include: snout–ventlength (SVL), head length (HL), eye–narialdistance (END), snout length (SNL), interor-bital distance (IOD), internarial distance(IND), eye diameter (ED), horizontal tym-panic annulus diameter (TAD), eye–tympa-num distance (ETD), head width (HW),upper arm length (UAL), forearm length(FAL), tibia length (TBL), femur length(FEL), tarsus length (TSL), pes length (PL),manus length (ML), Toe IV length (Toe4L),Finger I length (Fin1L), Finger III length(Fin3L), Finger I disc width (Fin1DW),Finger III disc width (Fin3DW), Toe IV discwidth (Toe4DW), and widths of penultimatephalanges of third finger (PpFin3) and thefourth toe (PpToe4). In the description,ranges are followed by mean 6 standarddeviation in parentheses. Specimens weredeposited at the Philippine National Museum(PNM) and Kansas University (KU). For thisdescription, we follow Kraus and Allison(2007) in maintaining the masculine forma-tions of species names within the genusPlatymantis.

RESULTS

Platymantis bayani sp. nov.(Figs. 2–4)

Holotype.—PNM 9501 (field no. CDS2989, formerly KU 309260; Fig. 2), an adultmale collected at 20:00 hr on 22 October 2007at 140 m elevation in Taft Forest (11u 4899.180 N, 125u 179 33.9360 E; WGS-84)Barangay San Rafael, Municipality of Taft,Eastern Samar Province, Samar Island, Phi-lippines, by Cameron D. Siler and Jason B.Fernandez.

Paratopotypes.—KU 309252–54, 309256,309258–59, 309261, 309263–64, 309266–67,and 309269, PNM 9515–22, eleven males andnine females, collected between 14 and 23October 2007.

Diagnosis.—Platymantis bayani can bedistinguished from other Philippine congeners

FIG. 1.—Known occurrence of Platymantis bayani ineastern Samar Island, Philippines (black dot). The insetshows the location of Samar Island (colored black) withinthe Philippines. The three provinces of Samar Island areindicated by dashed lines.

March 2009] HERPETOLOGICA 93

by the following combination of characters:body size 34.2–39.1 mm SVL for males, 44.3–49.8 mm SVL for females; digital disc ofFinger III 2.7–4.03 width of penultimatephalanx in males and 2.8–4.53 in females;digital disc of Toe IV 2.1–3.33 width ofpenultimate phalanx in males and 2.9–3.73

width in females (Fig. 3A,B); finger discs onlyslighter larger than toe discs; eye diameterlarge, 4.8–6.1 mm in adult males, and 5.4–6.8 mm in adult females; skin granular ondorsal surfaces; dorsal dermal tuberclessparsely distributed, salmon-colored; supra-tympanic fold moderately protuberant; tym-panum completely visible; dorsolateral stripesabsent; and a preference for terrestrialmicrohabitat in karst forest. Mensural anddiagnostic differences are provided in Ta-bles 1 and 2.

Comparisons.—The new species is assign-able to the P. guentheri Group (P. banahao, P.cornutus, P. diesmosi, P. guentheri, P. insula-tus, P. luzonensis, P. negrosensis, and P.rabori) based on the following combinationof characters: digital discs greatly expandedand larger in fingers than in toes, first fingeronly slightly shorter than second, proximalportions of digits with narrow dermal folds incross section, and subarticular tubercles largeand highly protuberant (Fig. 3).

FIG. 3.—Ventral views of hand (A) and foot (B) of male holotype of Platymantis bayani (PNM 9501), hand (C) andfoot (D) of male Platymantis insulatus (KU 300346), hand (E) and foot (F) of male Platymantis spelaeus (KU 300438),and hand (G) and foot (H) of male Platymantis paengi (KU 300209). Scale bar 5 2 mm.

FIG. 2.—Head of male holotype of Platymantis bayani(PNM 9501) in oblique view. Scale bar 5 5 mm.

94 HERPETOLOGICA [Vol. 65, No. 1

FIG. 4.—Photographs in life of Platymantis bayani paratopotypes exhibiting typical limestone-forest frog mottled colorpattern and salmon-colored dorsal tubercles; (A) KU 309254, female, SVL 5 49.3 mm, and (B) KU 309252, male, SVL5 34.7 mm. Photographs by C. Siler.


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From all members of the P. dorsalis Group(P. cagayensis, P. corrugatus, P. dorsalis, P.indeprensus, P. levigatus, P. mimulus, P.naomiae, P. paengi, P. pseudodorsalis, P. pyg-maeus, P. spelaeus, and P. taylori), Platyman-tis bayani is distinguished by the presence (vs.absence) of the following features: greatlyexpanded digital discs (Fig. 3); slightly largerfinger discs than toe discs; first finger shorterthan second finger (Fig. 3); and sparselydistributed, salmon-colored, dorsal dermaltubercles (vs. presence and dominant dorsalcolor in P. dorsalis, P. paengi, P. spelaeus, andP. taylori, and absence in P. cagayensis, P.corrugatus, P. indeprensus, P. levigatus, P.mimulus, P. naomiae, P. pseudodorsalis, andP. pygmaeus; Table 2). Platymantis bayani isfurther distinguished from all members of theP. dorsalis Group, except P. spelaeus, byhaving a much larger body size, and from allmembers except P. paengi and P. spelaeus, byits microhabitat preference (terrestrial sub-strates in limestone forest vs. leaf litter offorest floor and/or herb-layer vegetation in P.cagayensis, P. corrugatus, P. dorsalis, P. inde-prensus, P. mimulus, P. naomiae, P. pseudo-dorsalis, P. pygmaeus and P. taylori, or riverbanks and rocks in P. levigatus). From P.cagayensis, P. corrugatus, P. dorsalis, P. inde-prensus, P. mimulus, P. naomiae, P. pygmaeusand P. taylori, the new species is distinguishedby the absence (vs. presence) of dorsal foldsand ridges.

The new species is distinguished from all P.hazelae Group species (P. hazelae, P. isarog, P.lawtoni, P. montanus, P. panayensis, P. poli-llensis, P. sierramadrensis, P. subterrestris) byits larger body size; the first finger muchshorter than the second; narrow dermalphalanges on the digits; finger discs onlyslightly larger than toe disks; protuberant (vs.flattened) subarticular tubercles, and a pre-ferred microhabitat of terrestrial substrates inlimestone forest (vs. arboreal habitat, and/orshrub-layer vegetation).

In the following section, the new species iscompared with all species of the P. guentheriGroup. The character states for congeners canbe found in Tables 1 and 2.

The new species differs from all otherspecies of the P. guentheri Group, except forP. diesmosi, by its larger body size and greater

baya

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eye diameter. The new species differs from P.negrosensis, P. luzonensis, P. rabori, and P.guentheri by lacking rugose dorsal skin, dorsalfolds and ridges, and dorsolateral stripes. It isdistinguished from P. diesmosi, P. negrosensis,P. luzonensis, P. rabori, P. guentheri, and P.banahao by having a completely visibletympanum, dark brown to black spots on theflanks, sparsely distributed dorsal dermaltubercles that are salmon-colored, and apreference for terrestrial microhabitats inlimestone forest. Platymantis bayani is furtherdistinguished from P. negrosensis, P. luzonen-sis, and P. rabori by having a smooth and lessprominent supratympanic fold. From P. cor-nutus, P. negrosensis, P. luzonensis, and P.banahao, it differs in having slightly largerfinger discs than toe discs. The new speciesdiffers from P. diesmosi in having greatlyexpanded digital discs, and from P. cornutus,by having dark brown to black spots on theflanks, sparsely distributed, salmon-colored,dorsal dermal tubercles, a slightly prominentsupratympanic fold, well-defined limb bands,the absence of dorsolateral stripes, and apreference for terrestrial microhabitats inlimestone forest. From P. insulatus, the newspecies is distinguished by the presence ofdorsal dermal tubercles and a slightly prom-inent supratympanic fold. Finally, P. bayanidiffers from P. insulatus and P. banahao inhaving a distinct, transverse limb-bandingpattern.

Description of holotype.—A mature male;habitus robust; head broader than body, HL37.7% SVL, 91.2% HW; snout protrudingmoderately beyond lower jaw, snout tip comesto sharp point in dorsal and lateral aspects;

eyes large and protruding laterally, ED 15.6%SVL, 41.3% HL; lips slightly flared andswollen (Fig. 2); interorbital region flat; ED137.0% SNL, 1.73 END; pupil horizontallyelliptical; canthus rostralis laterally concave indorsal aspect; loreal region greatly concave,sloping ventrolaterally to labium; nostrilsoriented posterolaterally, protruding laterallyto sharp points from snout tip; END 2.33SNL; internarial region flat; tympanic annulusdistinct, its diameter 0.53 ED; supratympanicfold moderately protuberant, extending fromposterior corner of eye across dorsal margin oftympanum to supra-axillary region.

Tongue elongate, wider posteriorly, withdeep posterior notch and narrow anteriorattachment; choanae situated at anterolateraledge of palate, oval, widely separated bydistance three to four times greater thandiameter of single choana; dentigerous pro-cess of vomer distinct, with two teeth locatedon anterior 1/3 of process; dentigerous processangled slightly anterolaterally, approximatelyat 10–20u incline with closest (posterior)points separated by distance equal to 2.03diameter of single choana, their most distant(anterior) ends separated by distance equal to3.03 diameter of single choana; short vocalslits at posteroventral margin of mouth.

Skin of dorsal surfaces of trunk and headcoarsely granular, distinctly shagreen; irregular,moderately protuberant tubercles sparsely dis-tributed across dorsal surfaces of body and limbs;ventral surfaces of head smooth; lateral andventral surfaces of limbs finely granular withdorsal surfaces coarsely granular; flanks shagreen.

Manus length 58.0% pes length; tibia length108.8% pes length; tibia length 54.6% SVL;

TABLE 2.—Summary of mensural characters (in mm) and selected qualitative diagnostic characters (present, absent) inPlatymantis bayani and other Philippine limestone forest frog species. Sample size, body size, species group, and generalgeographical distribution (PAIC 5 Pleistocene Aggregate Island Complexes, sensu Brown and Diesmos, 2002) are

included for reference (SVL given as range over mean 6 standard deviation).

bayani sp. nov. (9m, 9f)insulatus(8m, 6f) spelaeus (4m, 3f) paengi (15m)

SVL (m) 34.2–39.1 34.3–38.0 37.1–44.3 27.7–34.3SVL (f) 44.3–49.8 40.2–42.9 53.5–57.5 N/ARange Samar Island Gigantes Islands SW Negros Island NW Panay IslandSpecies Group guentheri guentheri dorsalis dorsalisDigital disks Widely expanded Widely expanded Non-expanded Non-expandedDorsal skin rugosity 2 to granular 2 +, 2 +Dorsal dermal tubercles +, sparse 2 +, small +Flanks Spotted Spotted Immaculate Spotted


fingers slender, long; terminal discs greatlyexpanded (Fig. 3A) with pronounced distal,circummarginal groove; dorsal surfaces ofterminal phalanges with small, cutaneoussupra-articular flaps between ultimate andpenultimate phalanges; relative lengths offingers I 5 II , IV , III; subarticulartubercles prominent, convex, granular intexture; one subarticular tubercle belowDigits I and II, two tubercles under DigitsIII and IV; supernumerary tubercles distinct,round, prominent; present at bases of DigitsI–IV; inner metacarpal tubercle enlarged,elongate, crescent shaped; outer metacarpaltubercle partially divided into oval, slightlyraised preaxial tubercle of moderate size, andround postaxial tubercle equal in size topreaxial metacarpal tubercle; nuptial padsabsent; forearms slender, not hypertrophiedin males.

Tarsus smooth, lacking folds, flaps, ortubercles; terminal discs of toes greatlyexpanded, with circummarginal grooves; su-pra-articular cutaneous flaps above ultimate-penultimate phalangeal articulations; plantarsurface of foot smooth, with well-developed,prominently rounded subarticular tubercles(Fig. 3B); supernumerary tubercles absent;relative lengths of toes I , II , V , III ,IV; outer metatarsal tubercle small, round;inner metatarsal tubercle prominent, withsharp inner edge, length equal to distancefrom base of Toe I to first subarticulartubercle; toes with vestige of basal interpha-langeal web, nearly extending to the level ofthe proximal subarticular tubercles betweenToes III and IV.

Coloration of holotype in preservative.—Dominant dorsal color pattern on head andbody mottled with gray and silver blotches andspots interspersed throughout a dominantbackground of black and dark brown; dorsalsurfaces of limbs banded with alternatinggreen-brown and dark brown transverse bars;interorbital bar slightly darker than dominantdorsal color pattern; snout region slightlydarker than dorsum, and nearly uniformblack; wide black bar extending from snouttip, across canthus rostralis to anterior edge ofeye; a dark blotch covering anteroventral edgeof tympanum, ventral two thirds of tympanumlighter gray; temporal region dorsally mottled

dark brown and black; lips lighter gray withfaint vertical dark brown labial bars andspeckles, bordered anteriorly by lighter brownlower edge of canthus and subocular regions;transverse, dark brown bars on limbs similarin color to dorsum, coalescing into mottledpatterns on elbows and knees; four bars onforearms, four on thigh, four on tibia, three ontarsus; spotted flanks, ground color lightergrayish tan with dark brown blotches andspots, blending into the darker dorsal pattern(above) and the lighter ventral pattern (be-low); dorsal surfaces of manus, pes, and digitslight grayish tan with dark brown blotchesabove each joint.

Throat marbled light cream to tan withspeckled green-brown blotches; periphery ofthroat and lower lips lighter with anterioredge of lower lip possessing outline of lightcream spots; chest light cream with less densespeckled green-brown blotches; lighter ventralcoloration limited to small area on ventralsurface of forelimb near limb insertion;ventral surface of upper- and forearms darkercream with green-brown blotches; venter lightcream with nearly indistinct light green-brownblotches that decrease in size posteriorly untilblending with uniformly light cream groin;ventral surfaces of thigh uniformly mottledcream and green-brown; ventral surface oftibia and tarsus mottled green-brown, similarto forearms; palmar and plantar surfaces ofhands and feet dark green-brown with lightcream subarticular tubercles; iris pale grayabove and dark gray below pupil.

Coloration of holotype in life.—Dorsumand head mottled light green, dark green,and black (Fig. 4); interorbital bar distinct,black, and bordered anteriorly by lighter,green-brown border; dorsal surfaces of limbswith alternating dark brown and green trans-verse bars; dorsal surface of body with low,salmon-colored tubercles; series of darkbrown to black blotches and spots acrossflanks; pupil bordered by bright gold ciliaryring of iris, blending into the predominantmottled golden-brown and dark brown colorpattern of iris.

Measurements of holotype (mm).—SVL39.1; HL 14.7; HW 16.2; SNL 4.4; IND 3.6;IOD 3.2; ED 6.1; END 3.5; TAD 2.8; ETD1.0; UAL 7.4; FAL 9.0; FEL 20.1; TBL 21.3;


TSL 12.8; ML 11.4; PL 19.6; Fin1L 6.1;Fin3L 10.3; Toe4L 18.2; Fin3DW 1.8;Toe4DW 1.5; PpFin3 0.7; PpToe4 0.7.

Variation.—Summaries of univariate mor-phological variation in the series are presentedin Table 3. Dominant color pattern consis-tently mottled in all observed specimens;however, the degree of contrast between thedominant ground color and dark blotchesvaries among individuals, with two patterns ofcontrast observed. The first pattern consists ofdark brown to black blotches intermixed witha light cream to green ground color (KU309253–54, 309256, 309258–59, 309261, and309264). The second pattern was consists ofdark green blotches intermixed with the samelight cream to green ground color (KU309266, 309269, 309252, 309263, and309267).

Distribution.—Platymantis bayani is knownonly from the type locality (Fig. 1).

Etymology.—The name of the new speciesis derived from the Tagalog (Filipino) term‘‘bayani,’’ meaning ‘‘highly respected,’’ ‘‘hero-ic,’’ or ‘‘hero.’’ We are pleased to name this

new species for the late Walter C. Brown inrecognition of his numerous contributions toPhilippine herpetology and in acknowledg-ment of his lifelong commitment to the studyof biodiversity in the SW Pacific. Suggestedcommon name: Walter’s Limestone ForestFrog.

Ecology and natural history.—Platymantisbayani occurs in primary- and secondary-growth forest on karst substrate. The adver-tisement call of the species is unknown. Bothmales and females were found perched on topof limestone rocks within large rock forma-tions in the forest; they never were observedperching in trees or shrubs as do most speciesin the P. guentheri Group (R. M. Brown andGonzales, 2007).

Platymantis bayani was well camouflagedagainst the limestone (much like P. insulatus,P. speleaus, and P. paengi; Fig. 4), andcollectors found it difficult to locate andcapture individuals against the limestonesubstrate. When disturbed, the new speciesimmediately jumped back into the rocks andescaped into limestone crevices. No egg

TABLE 3.—Summary of univariate morphological variation among mensural characters in the type series of Platymantisbayani. See text for definitions of abbreviations.

Female n 5 9 Male n 5 9

SVL 44.3–49.8 (46.4 6 2.1) 34.2–39.1 (37.1 6 1.8)HL 16.2–18.8 (17.3 6 0.8) 14.1–15.5 (14.8 6 0.5)END 3.3–4.8 (4.1 6 0.5) 3.2–4.4 (3.9 6 0.4)SNL 3.8–6.1 (5.3 6 0.8) 4.0–5.9 (5.0 6 0.8)IOD 4.0–4.9 (4.3 6 0.3) 3.0–4.2 (3.5 6 0.3)IND 3.6–4.3 (3.9 6 0.2) 2.9–3.6 (3.2 6 0.2)ED 5.4–6.8 (6.1 6 0.4) 4.8–6.1 (5.5 6 0.5)TAD 2.3–3.3 (2.8 6 0.3) 1.9–2.8 (2.4 6 0.3)ETD 1.1–1.7 (1.4 6 0.2) 1.0–1.6 (1.3 6 0.2)HW 18.4–20.6 (19.7 6 0.7) 15.1–17.0 (15.9 6 0.6)UA 8.2–9.9 (9.1 6 0.6) 7.3–8.1 (7.6 6 0.3)FAL 12.0–13.1 (12.6 6 0.4) 9.8–10.6 (10.3 6 0.2)FL 23.7–26.2 (24.9 6 0.9) 19.6–20.7 (20.2 6 0.4)TBL 21.0–26.9 (25.8 6 1.8) 21.2–22.8 (21.7 6 0.5)TSL 14.7–15.9 (15.1 6 0.4) 12.0–13.2 (12.6 6 0.4)PL 22.9–25.3 (23.9 6 0.9) 18.7–20.3 (19.7 6 0.6)ML 13.5–14.8 (14.2 6 0.3) 11.2–12.3 (11.7 6 0.4)Toe4L 21.8–23.9 (22.8 6 0.6) 18.1–20.0 (18.8 6 0.6)Fin1L 7.2–8.4 (7.9 6 0.4) 6.1–7.1 (6.4 6 0.4)Fin3L 12.7–13.9 (13.2 6 0.4) 10.3–11.5 (10.8 6 0.4)Fin1DW 1.6–2.0 (1.8 6 0.1) 1.3–1.6 (1.4 6 0.1)Fin3DW 2.3–2.9 (2.7 6 0.2) 1.8–2.5 (2.1 6 0.2)Toe4DW 1.7–2.3 (2.0 6 0.2) 1.5–1.8 (1.6 6 0.1)PpFin3 0.6–0.8 (0.7 6 0.1) 0.5–0.7 (0.7 6 0.1)PpToe4 0.6–0.7 (0.6 6 0.1) 0.5–0.7 (0.6 6 0.1)


masses or juveniles were found; other speciesof Platymantis have direct development (W.C. Brown and Alcala, 1982a,b). Platymantisspelaeus has been observed by one of us(CDS) to deposit direct-developing eggs inlimestone crevices and the same is suspectedfor P. insulatus (R. M. Brown and Alcala,2000). We assume that this species also breedsin limestone crevices and small caves (R. M.Brown and Alcala, 2000; R. M. Brown et al.,2003).

Sympatric anuran species include Kalo-phrynus pleurostigma, Limnonectes magnus,Megophrys stejnegeri, Platymantis cf. dorsalis,P. cf. corrugatus, P. rabori, P. guentheri,Polypedates leucomystax, Philautus leitensis,Rhacophorus bimaculatus, R. appendiculatus,R. pardalis, Oreophryne sp., Rana grandocula,R. albotuberculata, and Occidozyga laevis.

DISCUSSION

With the new species described here, a totalof 28 Philippine species of Platymantis arenow recognized. The species total for thecountry has increased dramatically over thelast several decades; seven species wererecognized between 1950 and 1970 (Inger,1954; W. C. Brown and Alcala, 1970a,b) and12 by the late 1980’s (W. C. Brown and Alcala,1982a). An additional dozen species weredescribed in the late 1990’s (W. C. Brown etal., 1997a,b,c, 1999), several more specieshave been described recently (R. M. Brownand Gonzales, 2007; Siler et al., 2007), andnumerous unnamed species await description(Brown, 2004; Brown et al., 2008).

Within the Philippine radiation, high levelsof cryptic diversity (Alcala et al., 1998; W. C.Brown et al., 1997b, 1999) has only recentlybeen uncovered (Brown, 2004; Brown et al.,2008) due to attention on advertisement-calldiversity, microhabitat preferences, and genesequence data

Platymantis bayani is the fourth Philippinelimestone frog to be described, the thirdterrestrial member of the P. guentheri Group(the others being P. insulatus and P. diesmosi),and the only limestone species known to occurin the Mindanao Faunal Region. Platymantisspelaeus from SW Negros Island, P. insulatusfrom the Gigantes Islands, and P. paengi fromNW Panay Island are the three previously

known karst-forest frogs found in the Visayan(Central) Philippine Islands. All four speciesseem to be obligate inhabitants of limestoneforests. Although potentially restricted ingeographic ranges, the species are quitecommon in the rainy season, but can seemto be rare in times of high temperature andaridity when we assume they retreat intosubterranean limestone caves and crevices (R.M. Brown and Alcala, 2000; R. M. Brown etal., 2003).

The recent discoveries of new species ofPhilippine frogs and gekkos with preferencesfor limestone forests (Siler et al., 2007;Roesler et al., 2006) highlight the degree towhich this imperiled habitat type (Clements etal., 2006) is poorly known. Future surveyefforts should make a point to target limestoneforest communities. Our general impression isthat these habitat types have been overlookedin the past because surrounding forests aredry, stunted, difficult to sample, and viewedby field biologists as non-productive.

The heavily mottled, predominant colorpattern of Platymantis bayani resembles P.spelaeus, P. insulatus, and P. paengi. Theconspicuous similarity of color patterns amongthese taxa suggests to us the possibility ofconvergent selection on color pattern inlimestone habitats. It is unclear as to whichspecies P. bayani is most closely related, butphenotypic similarity with P. insulatus isstriking.

The new species was quite common at thetype locality, and it may posses a widergeographical distribution, possibly throughoutthe northern islands of the Mindanao FaunalRegion (Samar, Leyte, Biliran, Mindanao,Dinagat, Siargao, and smaller associatedislands). It is possible that a primary limitingfactor to the range of P. bayani is the presenceof relatively undisturbed limestone formationsand forest cover. We note that both P.speleaus and P. insulatus persist quite well,despite the near complete removal of forest onthe Gigante Island group (P. insulatus) andsouthern Negros Island (P. spelaeus). Inten-sive survey efforts will be required to fullydocument the distribution of the new speciesand arrive at a reasonable assessment of itsconservation status. At present, we considerthe status of the new species to be ‘‘data


deficient’’ pending the collection of muchneeded data on its distribution and habitatrequirements.

Acknowledgments.—We thank the Protected Areas andWildlife Bureau (PAWB) of the Philippine Department ofEnvironment and Natural Resources (DENR) for facili-tating collecting and export permits necessary for this andrelated studies, wherein we are particularly grateful to M.Lim, C. Custodio, and A. Tagtag. We also thank the ArnieViojan and the Regional Office of the Protected Areas,Wildlife, and Coastal Zones Management Services(PAWCZMS), and Director Manolito D. Ragub, EirishMate, and the headquarters of the Samar Island NaturalParks (SINP), whose logistical support and guidance wereinstrumental throughout our visit to Samar. JasonFernandez assisted in the field. Financial support forfieldwork was provided by a Panorama Fund grant toCDS from The University of Kansas Natural HistoryMuseum and Biodiversity Institute, a Madison and LilaSelf Fellowship to CDS from the University of Kansas,NSF EF-0334952 to L. Trueb and R. Brown and DEB0743491 to RMB. For the loans of specimens (museumabbreviations follow Leviton et al., 1985), we thank J.Vindum and A. Leviton (CAS), R. Sison and A. C.Diesmos (PNM), J. Ferner (CMNH), A. Resetar and H.Voris (FMNH), R. Crombie (USNM), and T. LaDuc(TNHC). Critical reviews of the manuscript wereprovided by L. Trueb and J. Esselstyn. We thank MichaelGarfield for illustrating the holotype for Figure 2. CDSthanks the Philippine-American Education Foundation,A.C. Alcala and family, and M. Diesmos for theircontinued support, as well as CAS’s Stearns Fellowshipfor funding a recent visit to examine comparative material.Finally, we owe a debt of gratitude to the late Walter C.Brown who alerted RMB to the presence of this newspecies on Samar more than 10 yr before we were able tofind it. Once again, Walter was right.

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ZWEIFEL, R. G. 1975. Frogs of the genus Platymantis(Ranidae) from New Britain. American MuseumNovitates 7582:1–7.

.Accepted: 13 February 2009

.Associate Editor: Michael Harvey

APPENDIX I

Specimens Examined

All specimens examined are from the Philippines.Numbers in parentheses indicate the number of speci-mens examined for each species. Platymantis bana-hao.—(6) LUZON ISLAND, QUEZON PROVINCE, Munic-ipality of Tayabas, Baragany Lalo, Mt. Banahao: TNHC61968–71, PNM 9248–49. Platymantis cagayanensis.—(9) LUZON ISLAND, CAGAYAN PROVINCE, ‘‘Tagat ForestReserve near Santa Praxedes Town:’’ PNM 7564, 7578,7496–99, 7506, 7608, 7526. Platymantis cornutus.—(3)LUZON ISLAND, KALINGA PROVINCE, Municipality ofBalbalan, Barangay Balbalan: CAS 231498, 231501,CMNH 8128. Platymantis corrugatus.—(22) CAMI-GUIN ISLAND, CAMIGUIN PROVINCE, Municipality ofGuinsiliban, Barangay Cabuan: KU 300351, 300355;POLILLO ISLAND, QUEZON PROVINCE, Municipality ofPolillo, Barangay Pinaglubayan, 62 m elevation, 14u 4599.30 N, 121u 589 5.520 E: KU 300350, 300352–54;NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Munic-ipality of Valencia, Barangay Bongbong, Camp Lookout,Cuernos de Negros Mt. Range, Mt. Talinis, 500 melevation: TNHC 61972–87. Platymantis diesmosi.—(10) LUZON ISLAND, ALBAY PROVINCE, Municipality ofTiwi, Barangay Banhaw: PNM 8499 (Holotype), 8500–1(Paratypes), TNHC 62040–42 (Paratypes), UPLB-MNH16, 21–23 (Paratypes). Platymantis dorsalis.—(22)


NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Munic-ipality of Valencia, Barangay Bongbong, Sitio Tagaytay,Mt. Talinis, 1150 m elevation, 9u 159 360 N, 123u 129 1960E: KU 300356–300377. Platymantis guentheri.—(19)DINAGAT ISLAND, SURIGAO PROVINCE, Municipality ofLoreto, Barangay Esperanza: KU 306321–23, 306325;SAMAR ISLAND, EASTERN SAMAR PROVINCE, Municipal-ity of Taft, Barangay San Rafael: KU 309185, 309189,309191, 309195, 309203, 309217–19, 309221, 309228–30,309232, 309236, 309238. Platymantis hazelae.—(13)NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Munic-ipality of Valencia, Barangay Bongbong, Sitio Tagaytay,Mt. Talinis, 1150 m elevation: KU 300403–15. Platy-mantis indeprensus.—(8) LUZON ISLAND, QUEZON

PROVINCE, Municipality of Tayabas, Barangay Lalo, Mt.Banahao: TNHC 061956–60; PNM 9257–59. Platyman-tis insulatus.—(18) SOUTH GIGANTE ISLAND,ILOILO PROVINCE, Municipality of Carles, Barangay Gabi:CAS 117441 (Holotype), 119967–69 (Paratypes); KU300338–44, 300346, 309088–89; NORTH GIGANTEISLAND, ILOILO PROVINCE, Municipality of Carles,Barangay Granada: KU 300345, 300347–49. Platymantisisarog.—(7) LUZON ISLAND, CAMARINE SUR PROVINCE,Municipality of Naga City, Barangay Panicuason, Mt.Isarog National Park, Mt. Isarog: TNHC 61961–67.Platymantis lawtoni.—(1) TABLAS ISLAND, ROMBLON

PROVINCE, Municipality of San Agustin, Mt. Progreso:CAS 135733. Platymantis levigatus.—(15) SIBUYANISLAND, ROMBLON PROVINCE, Municipality of Magdi-wang, Barangay Talaba, Mt. Guiting-Guiting NaturalPark, 0 m elevation: KU 300416–30. Platymantis luzo-nensis.—(27) LUZON ISLAND, LAGUNA PROVINCE,Municipality of Los Banos, Barangay Batong Malake,Mt. Makiling: CAS 196364, 196369–70, 200404–08,210544–45 (Paratypes); CAMARINES SUR PROVINCE, Munic-ipality of Naga City, Mt. Isarog: FMNH 251643–44;TNHC 62004–09, 62012–13, 62020–24; POLILLO IS-LAND, QUEZON PROVINCE, Municipality of Polillo, Bar-angay Pinaglubayan: KU 305541–42. Platymantis mimu-lus.—(12) LUZON ISLAND, LAGUNA PROVINCE,Municipality of Los Banos, Barangay Batong Malake,Mt. Makiling: TNHC 54930–31; PNM 9260–69. Platy-mantis montana.—(13) LUZON ISLAND, QUEZON

PROVINCE, Municipality of Tayabas, Barangay Lalo, Mt.Banahao: TNHC 62149–58; CAS 200998–1000. Platy-mantis naomiae.—(3) LUZON ISLAND, QUEZON PROV-

INCE, Municipality of Tayabas, Baragany Lalo, Mt.Banahao: TNHC 62169–71. Platymantis negrosen-sis.—(7) NEGROS ISLAND, NEGROS ORIENTAL PROV-

INCE, Cuernos de Negros Mountain Range, Mt. Talinis,750 m elevation: KU 300439–45. Platymantis paengi.—(15) PANAY ISLAND, ANTIQUE PROVINCE, Municipality ofPandan, Barangay Duyong: PNM 9239 (Holotype), 9240–43 (Paratopotypes), KU 300206–13 (Paratopotypes),300204–05 (Paratypes). Platymantis panayensis.—(2)PANAY ISLAND, AKLAN PROVINCE, Municipality ofNabas: CAS 137641–42. Platymantis pseudodorsa-lis.—(4) LUZON ISLAND, QUEZON PROVINCE, Munici-pality of Tayabas, Baragany Lalo, Mt. Banahao: KU207455–57, 207459 (Paratypes). Platymantis pyg-maeus.—(5) LUZON ISLAND, CAGAYAN PROVINCE,Municipality of Calveria, Barangay Mabnang, MabnangFalls: PNM 7523, 9528–31. Platymantis rabori.—(12)MINDANAO ISLAND, NEW BATAAN PROVINCE, Mt.Puting Bato: CMNH 2305, 2350; DAVAO CITY PROVINCE,Municipality of Toril, Barangay Baracatan, Sitio UpperBaracatan: CMNH 1462; Municipality of Calinan, BarangayMalagos: PNM 9504–05; SAMAR ISLAND, EASTERN SAMAR

PROVINCE, Municipality of Taft, Barangay San Rafael: KU309121–27. Platymantis sierramadrensis.—(6) LUZONISLAND, AURORA PROVINCE, Municipality of San Luis;Dipiningan branch of the Kobatangan River drainage; 15u409 120 N, 121u 209 480 E: CMNH 5678–79, 5904; ISABELA

PROVINCE, Municipality ofPalanan, Barangay Didian, SitioNatap Dukan, Northern Sierra Madre National Park, 16u579 55.80 N, 122u 249 13.80 E: CAS 204739–41. Platymantisspelaeus.—(7) NEGROS ISLAND, NEGROS ORIENTAL

PROVINCE, Municipality of Basay, Tiyabanan Barrio: CAS153477–78, 153482 (Paratypes); NEGROS OCCIDENTAL PROV-

INCE, Municipality of Cauayan, Sitio Banso, BarangayCamalandaan, 320 m elevation: KU 300435–38. Platyman-tis subterrestris.—(3) LUZON ISLAND, MOUNTAIN

PROVINCE, Mt Data: CAS 204319–204321. Platymantistaylori.—(4) LUZON ISLAND, ISABELA PROVINCE, Munic-ipality of Palanan, Barangay Didian, Sitio Natapdukan: CAS207443–207446 (Paratypes).


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