A NEW SPECIES OF PLATYMANTIS (AMPHIBIA: ANURA…€¦ · A NEW SPECIES OF PLATYMANTIS (AMPHIBIA:...

A NEW SPECIES OF PLATYMANTIS (AMPHIBIA: ANURA: RANIDAE)FROM PANAY ISLAND, PHILIPPINES

CAMERON D. SILER1,4, CHARLES W. LINKEM1, ARVIN C. DIESMOS

2, AND ANGEL C. ALCALA3

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: We describe a new species of forest frog (genus Platymantis) from 180–300 m above sea levelon Mt. Lihidan in the northwestern part of Panay Island, Philippines. It is assigned to the Platymantisdorsalis species group and is distinguished from congeners by external morphology, various spectral andtemporal components of the advertisement call, and a preference for terrestrial, limestone microhabitat.Unique morphological characters include a moderately large body (27.7–34.3 mm SVL for 15 males), slightlyexpanded terminal finger and toe discs, distinctly rugose dorsal and lateral skin, pronounced supratympanicfold, protuberant rictal tubercle cluster, hidden dorsal and posterior edges of tympanum, and uniquecoloration.

Key words: Advertisement calls; Biodiversity; Cryptic species; Endemism; Limestone frogs; Philippines;Platymantis dorsalis species group; SW Pacific

ANURANS of the genus Platymantis have twomajor centers of diversity, one in the Solomon(Brown, 1952), Bismarck (Allison and Kraus,2001; Foufopoulos and Brown, 2004; Zweifel,1960, 1975), and Admiralty archipelagos (21species; Allison, 1996), and the second in thePhilippines (27 species; Alcala and Brown,1998, 1999; Brown et al., 1997a; Brown andGonzales, in press). In addition, severalspecies occur outside of these major archipel-agos. One species occurs in Palau (Crombieand Pregill, 1999), seven in New Guinea(Allison, 1996; Gunther, 1999, 2005; Zweifel,1969), two to four in eastern Indonesia (Edgarand Lilley, 1993; Menzies, 1982a,b), and twoin Fiji (Gorham, 1965; Morrison, 2003).

In October 2004 and November 2005, weconducted herpetological field surveys in thelower-elevation forest of Mt. Lihidan in NWPanay Island (Fig. 1). Two individuals of anundescribed species of Platymantis, alongwith a recording of its advertisement call,were collected in 2004 at an elevation of300 m. Thirteen additional specimens andfour additional acoustic recordings wereobtained at lower elevations in 2005. In thispaper we describe the new species and reporton its natural history, ecology, habitat, andunusual advertisement call.

MATERIALS AND METHODS

We recorded morphometric data fromalcohol-preserved specimens that were origi-nally fixed in 10% formalin (Appendix 1). Sexwas determined by gonadal inspection whenpossible, and measurements were taken with

Herpetologica herp-63-03-11.3d 14/7/07 14:47:43 351 Cust # 06-067

4 CORRESPONDENCE: e-mail, [email protected]

FIG. 1.—Known distribution of Platymantis paengi onMt. Lihidan, NW Panay Island, Philippines. The insetshows the location of Panay Island (colored black) withinthe Philippines. The type locality (Mt. Lihidan, Munici-pality of Pandan, Antique Province) is indicated by a blackdot. The four provinces of Panay Island are indicated bydashed lines.

Herpetologica, 63(3), 2007, 351–364

E 2007 by The Herpetologists’ League, Inc.

351

digital calipers to the nearest 0.1 mm. Tominimize inter-observer bias and othersources of potential error (Hayek et al.,2001; Lee, 1982), all measurements werescored by CDS.

Characters measured include: snout–ventlength (SVL, from snout tip to posteriormargin of vent), head length (from tip ofsnout to posterior margin of jaw articulation),eye–narial distance (from anterior margin ofeye to posterior margin of nares), snout length(from anterior margin of eye to tip of snout),interorbital distance (between orbits at theirmidpoint), internarial distance (from dorsalaspect between most laterally distal edges ofnares), eye diameter (at widest point), hori-zontal tympanic annulus diameter (at widestpoint), eye–tympanum distance (from poste-rior margin of eye to anterior margin oftympanum), head width (widest measure ofhead width measured at jaw articulations),upper arm length (from margin of bodyinsertion to elbow tip), forearm length (fromelbow to proximal end of outer metacarpaltubercle), tibia length (from knee to heel offlexed leg), femur length (from cloaca to theouter surface of the flexed knee), tarsus length(from knee to proximal edge of metatarsaltubercle), pes length (from proximal edge ofouter metatarsal tubercle to tip of Toe IV),manus length (from tip of Finger III toproximal margin of the outer metacarpaltubercle), Toe IV length (from distal marginof metatarsal tubercle to Toe IV tip), Finger Ilength (from proximal margin of palmartubercle to Finger I tip), Finger III length(from distal margin of outer metacarpaltubercle to Finger III tip), Finger I disc width(at widest point), Finger III disc width (atwidest point), Toe IV disc width (at widestpoint), and widths of penultimate phalanges ofthird finger and the fourth toe (at midpoint ofpenultimate phalanges). In the description,ranges are followed by mean 6 SD inparentheses.

Advertisement calls were recorded witha Sony2 PCM-M1 Professional Digital AudioRecorder with a Sennheiser2 ME80 condens-er microphone (equipped with K3U powermodule). Calls were recorded by CDS atdistances of 1–4 m from calling frogs; ambi-ent, substrate, and cloacal temperatures were

collected immediately after calls were re-corded. Calls were digitized using SoundeditE(Macromedia, 1995) and analyzed using Ra-venE (Charif et al., 2004) software. Temper-ature corrections were not deemed necessarybecause ambient temperature was the samefor all recordings, and all frogs were recordedin close proximity to one another. Weexamined oscillograms (waveforms), audio-spectrograms (sonograms), and results of theFast Fourier Transformation (FFT; powerspectrum) for a series of spectral and temporalcall characteristics (Brown et al., 2003;Diesmos et al., 2002; Foufopoulos and Brown,2004). Definitions of advertisement call ter-minology follow Ron (2005). For this de-scription, we follow Kraus and Allison (inpress) in maintaining the masculine forma-tions of species’ names within the genusPlatymantis.

RESULTS

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

Holotype.—PNM 9239 (field no. CDS1585; Fig. 2), an adult male collected at20:15 h on 26 November 2005 at 180 melevation in an area known locally as ‘‘Mt.Lihidan’’ (11u 249 52.740 N, 122u 069 16.740 E;WGS-84) Barangay Duyong, Municipality ofPandan, Antique Province, Panay Island,Philippines, by Cameron D. Siler and CharlesW. Linkem.

Paratopotypes.—KU 300206–13, PNM9240–43, 12 males, collected on 25 and 26November 2005.

Paratypes.—KU 300204–05, two malescollected between 2000 and 2300 h on 16May 2004 at the same general locality asholotype and paratopotypes at 300 m in thefoothills of ‘‘Mt. Lihidan’’ (11u 459 39.30 N,122u 029 28.260 E; WGS-84) by Cameron D.Siler, L. Averia, and A. Ong.

Diagnosis.—Platymantis paengi can bedistinguished from other Philippine congenersby the following combination of characters:body moderately large in males (27.7–34.3 mm SVL, n 5 15); digital discs of fingers(1.3–2.0 [1.6 6 0.2] 3 width of penultimatephalange) and toes (1.4–2.5 [2.0 6 0.3] 3width of penultimate phalange) slightly ex-


352 HERPETOLOGICA [Vol. 63, No. 3

panded (Fig. 3); skin rugose on dorsal andventral surfaces; rictal tubercle cluster pro-tuberant; supratympanic fold rugose and pro-tuberant; tympanic annulus large (9% SVL);and advertisement call consisting of frequencysweeps (change in call frequency over time)and amplitude modulation (varying noteamplitudes across a single call) (Table 1).

Comparisons.—Three species groups with-in Philippine Platymantis were proposed byBrown et al. (1997a,c) on the basis of externalmorphology: the P. dorsalis Group, the P.hazelae Group, and the P. guentheri Group.The new species is a member of the P. dorsalisGroup (P. dorsalis, P. corrugatus, P. levigatus,P. cagayensis, P. taylori, P. pseudodorsalis, P.indeprensis, P. spelaeus, P. mimulus, P.naomiae, and P. pygmaeus), with the followingcombination of characters: digital discs onlyslightly to moderately expanded and smaller infingers than in toes, Fingers II and III

approximately equal in length, proximal por-tions of digits round in cross section, largesubarticular tubercles pointed and highlyprotuberant, and a terrestrial microhabitatpreference.

Platymantis paengi and P. spelaeus are theonly species of the P. dorsalis Group that arefound exclusively in limestone forest. Theremaining species occur on soil dominatedforest floors (P. pygmaeus, P. naomiae, and P.mimulus, P. pseudodorsalis, P. dorsalis) and/orshrub layer vegetation (P. cagayensis and P.indeprensis), and banks of fast-flowing streams(P. levigatus). In addition, Platymantis paengiproduces a unique advertisement call, consist-ing of a series of slight frequency modulatednotes, with the first note being the longest(Fig. 5E). The remaining species of the P.dorsalis Group produce single pulse calls (P.pygmaeus), complex, multi-syllable calls (P.naomiae, P. mimulus, P. corrugatus, P. ca-gayensis, P. taylori, P. indeprensis, and P.spelaeus), or pure-tone, frequency sweep calls(P. dorsalis, P. levigatus, and P. pseudodorsalis)


FIG. 3.—Ventral views of hand (A) and foot (C) of malePlatymantis paengi (KU 300209), and of hand (B) and foot(D) of male Platymantis dorsalis (KU 300356).

FIG. 2.—Head of male holotype of Platymantis paengi(PNM 9239) in dorsal and lateral views. Illustrationby CDS.

September 2007] HERPETOLOGICA 353


FIG. 4.—Photographs in life of Platymantis paengi paratopotypes exhibiting two color patterns: (A) the completelymottled phase (PNM 9241), male, SVL 34.3 mm, Photo: CWL; and (B) the solid rusty-orange dorsal phase (KU300206), male, SVL 34.0 mm, Photo: CDS.



TABLE1.—

Summaryof

qualitativediagno

sticcharacters

(present,absen

t)in

Platym

antispa

engi

andmalespecim

ensof

othe

rmem

bers

oftheP.

dorsalisspeciesgrou

p.Sample

size,b

odysize

ofmales,and

gene

ralg

eographicaldistribu

tion

(PAIC

5Pleistocene

Aggregate

Island

Com

plexes,sen

suBrownandDiesm

os,2

002)

areinclud

edforreference

(SVLgivenas

rang

eover

mean6

standard

deviation;

TAD/SVLgivenas

percen

tage

over

mean6

standard

deviation).P

.mim

ula,P.

naom

iae,andP.

pygm

aeawereno

tinclud

edbe

causeof

theirsm

allerbo

dysize,andP.

spelaeaisinclud

eddu

eto

microhabitatsimilarity.

paengi

(15)

dorsalis(13)

corrug

ata(17)

levigata

(14)

cagayensis(9)

pseudo

dorsalis(4)

taylori(4)

indeprensa

(7)

spelaea(4)

SVL(m

m)

27.7–34.3

(31.86

2.0)

24.9–35.4

(28.36

2.6)

27.2–33.9

(28.36

2.6)

24.1–33.2

(28.56

3.1)

24.0–28.0

(26.56

1.4)

23.1–29.8

(27.76

3.1)

27.1–33.0

(29.06

2.3)

25.5–27.9

(26.66

0.8)

37.1–44.3

(41.36

3.2)

TAD/SVL

8.0–10.0

(0.096

0.01)

8.0–10.0

(0.076

0.01)

5.0–7.0

(0.066

0.01)

4.0–7.0

(0.056

0.01)

5.0–7.0

(0.076

0.01)

5.0–8.0

(0.076

0.01)

7.0

(0.076

0.00)

6.0–8.0

(0.076

0.01)

6.0–8.0

(0.076

0.01)

Range

NW

Panay

Island

Philip

pine

sPhilip

pine

sRom

blon

Island

Group

Luzon

PAIC

Luzon

PAIC

Luzon

PAIC

Luzon

PAIC

SWNegros

Island

Digital

disk

expansion

Present

Absen

tAbsen

tPresent

Absen

tAbsen

tAbsen

tAbsen

tPresent

Rictaltube

rcle

clustersize

Present,large

Present,often1

tube

rcle

Absen

tPresent,1–2

tube

rcles

Present

orabsent

Present,sm

all

Absen

tto

very

small

Present,sm

all

Present,often1

tube

rcle

Dorsalskin

Rug

osity

Present

Present

orabsent

Absen

tto

presen

t,less

prom

inen

tAbsen

tAbsen

tPresent

orabsent

Absen

tPresent

orabsent

Present

orAbsen

tDorsalde

rmal

Tub

ercles

Present

Present

orabsent

Absen

tAbsen

tAbsen

tAbsen

tPresent,sm

all,

infreq

uent

Absen

tPresent,sm

all

Dorsalfolds/ridg

esPresent,irregu

lar

Present,

irregu

lar

Present,2

continuo

us,

irregu

lar

Absen

tpresen

t,irregu

lar,lowAbsen

tPresent,sm

all,

infreq

uent

Present,sm

all

Absen

t

Tym

panu

mhidd

enDorsal/p

osterior

edge

No

No

No

No

No

Dorsaled

geDorsaled

geNoto

dorsal

edge

Supratym

panicfold

Highlyrugose,

prom

inen

tSm

ooth,less

prom

inen

tSm

ooth,less

prom

inen

tLow

toindistinct

Lessprom

inen

tMod

erately

prom

inen

tMod

erately

prom

inen

tSm

ooth,less

prom

inen

tLessprom

inen

t

Palmar

metacarpal

tube

rcle

size/

separation

Large/present

Large/present

toabsent

Large/present

Axial

very

small/

presen

tLarge/present

Large/present

Large/present

Large/present

Large/present

Microhabitat

preferen

ceKarst,lim

estone

Forest

Forest

River

bank

,rocks

Forest

Forest

Forest

Forest

Karst,lim

estone

Advertisemen

tCall

Frequ

ency

sweep/

amplitud

emod

ulation

Pure-tone

freq

uency

sweep

Com

plex,

multi-syllable

Pure-tone

freq

uency

sweep

Com

plex,

multi-syllablePure-tone

freq

uency

sweep

Com

plex,

multi-syllable

Com

plex,

multi-syllableC

omplex,

multi-syllable


The new species differs from P. mimulus, P.naomiae, and P. pygmaeus by its larger bodysize (male P. paengi, SVL 5 27.7–34.3 mm,vs. 14.1–15.7 mm in P. pygmaeus [Alcala etal., 1998], 20.3–23.9 mm in P. naomiae [Alcalaet al., 1998], and 19.0–24.5 mm in P. mimulus[Brown et al., 1997b]), and microhabitatpreference (limestone forest vs. soil dominat-ed forest floor, and/or herb-layer vegetation).From larger members of the P. dorsalisGroup, Platymantis. paengi is distinguishedfrom P. pseudodorsalis by its slightly expanded(vs. narrow) terminal discs; a large, highlyprotuberant (vs. small, and less pronounced)cluster of rictal tubercles; having the dorsaland posterior edges of the tympanic annuluscovered (vs. completely distinct from) by thesupratympanic fold; the presence (vs. ab-sence) of dorsal folds and ridges; and the

presence (vs. absence) of dorsal dermaltubercles. From P. cagayensis and P. inde-prensis, P. paengi is distinguished by thepresence (vs. absence) of heavily tuberculatedorsal and lateral skin, the presence of a large(vs. absence or small) rictal tubercle cluster,and the presence (vs. absence) of a protuber-ant, rugose supratympanic fold.

Platymantis paengi is distinguished from P.spelaeus by its smaller size (male P. paengi,SVL 5 27.7–34.3 mm vs. 37.1–44.3 mm in P.spelaeus); a large, highly protuberant rictaltubercle cluster (vs. small, and often consist-ing of a single tubercle); the presence of largerand more abundant (vs. smaller and lessabundant) dorsal dermal tubercles; and thepresence (vs. absence) of dorsal folds andridges. The new species differs from P.corrugatus by the absence (vs. presence) ofa dark, pigmented bar covering the lateralportion of the head; numerous, small super-numerary tubercles across the ventral surfaceof the foot; and two to five long, discontinuousfolds down the dorsal surface. From P.levigatus, the new species differs by havingthe dorsal and posterior edges of the tympanicannulus covered (vs. completely distinct from)by the supratympanic fold; a supratympanicfold that extends fully to (vs. terminating wellbefore) the supra-axillary region; a sharplyangular (vs. rounded) canthus rostralis; andhighly rugose (vs. smooth) dorsal and lateralskin. The new species is distinguished from P.dorsalis by having slightly expanded (vs.narrow) terminal discs (Fig. 3); a large, highlyprotuberant (vs. small, and less pronounced)cluster of rictal tubercles; dorsal and posterioredges of the tympanic annulus covered by (vs.completely distinct from) the supratympanicfold; highly tuberculate (vs. smooth to slightlytuberculate) dorsal and lateral skin; a distinct(vs. absent or less well defined) bandingpattern on the limbs.

Description of holotype.—A mature male;habitus robust; head only slightly broader thanbody, head length 37.1% SVL; head length87.1% head width; snout protruding moder-ately beyond lower jaw, snout tip rounded indorsal and lateral aspect (Fig. 2); eyes pro-truding laterally; lips moderately flared andswollen; interorbital region flat; eye diameter80.5% snout length, 1.3 3 eye-nares distance;


FIG. 5.—The advertisement call of Platymantis paengi(Holotype, PNM 9239) recorded at ambient temperatureof 25.0 C, perch temperature of 25.0 C, and cloacaltemperature of 26.2 C on 26 November 2005: (A) powerspectrum (Fast Fourier Transformation; relative intensityin dB vs. frequency in kHz), (B) an oscillogram (relativeamplitude vs. time in s) of the final three calls from onecall group, (C) an expanded waveform (relative amplitudevs. time in ms) of a single call within a call group, (D) anaudiospectrogram (frequency in kHz vs. time in s), and(E) an expanded audiospectrogram of three notes showingslight frequency modulation over time.


pupil horizontally elliptical; canthus rostralislaterally concave in dorsal aspect; loreal regionconcave, sloping ventrolaterally to labium;nostrils oriented posterolaterally; eye-naresdistance 2.8 3 nostril–tip of snout distance;internarial region slightly concave; tympanicannulus distinct, its diameter 0.78 3 eyediameter; dorsal and posterodorsal margin oftympanic annulus concealed by supratympa-nic fold; fold strongly protuberant, extendingfrom posterior corner of eye across dorsalmargin of tympanum to supra-axillary region.

Tongue elongate, wider posteriorly, withdeep posterior notch and narrow anteriorattachment; choanae situated at anterolateraledge of palate, round, widely separated bydistance four to five times greater thandiameter of single choana; dentigerous pro-cess of vomer distinct, with six teeth; denti-gerous process angled slightly anterolaterally,approximately at 20–30u incline with closest(posterior) points separated by distance equalto 1.5 times diameter of single choana, theirmost distant (anterior) ends separated bydistance equal to three times diameter ofsingle choana; short vocal slits at poster-oventral margin of mouth.

Skin of dorsal surfaces of trunk and headdistinctly shagreen, bearing heterogeneoustubercles (tiny and barely perceptible to largeand highly pronounced) and highly pro-nounced dorsal ridges; ridges irregular inlength, none spanning entire body length,and concentrated in suprascapular and sacralregion of dorsum; ventral surfaces of headsmooth; lateral and ventral surfaces of limbsfinely granular with dorsal surfaces moderate-ly tuberculate; flanks shagreen.

Manus length 50.5% pes length; tibia length99.5% pes length; tibia length 55.6% SVL;fingers slender, long; terminal discs slightly tomoderately expanded (Fig. 3A) with pro-nounced distal, circummarginal groove; dorsalsurfaces of terminal phalanges with small,cutaneous supra-articular flaps between ulti-mate and penultimate phalanges; relativelengths of fingers: II 5 IV , I , III;subarticular tubercles prominent, convex,velvety in texture; one subarticular tuberclebelow Digits I and II, two tubercles underDigits III and IV; supernumerary tuberclesdistinct, round, prominent; present at base of

Digits I–IV; thenar (inner metacarpal) tuber-cle enlarged, elongate, ovoid; palmar (outermetacarpal) tubercle completely divided intofollowing: preaxial tubercle large, oval, prom-inently raised; postaxial tubercle oval, half thesize of preaxial metacarpal tubercle; nuptialpads absent, forearms slender.

Tarsus smooth, lacking folds, flaps, ortubercles; terminal discs of toes moderatelyexpanded, with circummarginal grooves; su-pra-articular cutaneous flaps above ultimate-penultimate phalangeal articulations; plantarsurfaces of foot smooth, with well-developed,prominently rounded to pointed subarticulartubercles (Fig. 3C); relative lengths of toes: I, II , V , III , IV; outer metatarsaltubercle small, round, pointed; inner meta-tarsal tubercle prominent, with sharp plantaredge, length equal to distance from base ofToe I to first subarticular tubercle; toes barelywebbed basally (not extending to level of theproximal subarticular tubercles); cloacal re-gion glandular, lacking tubercles or supracloa-cal flaps.

Coloration of holotype in preservative.—Dominant dorsal color on head and bodynearly uniform dark brown with distinct, palegray patches; dorsal surfaces of limbs mediumbrown with transverse, dark brown bars;interorbital bar faint, slightly paler, borderedposteriorly by dark brown and anteriorly byslightly paler snout; wide black bar extendingfrom nares through canthal, preocular, posto-cular, and tympanic regions, wrapping ven-trally around posterior edge of tympanicannulus where it expands to cover anteroven-tral edge of tympanum; dark colorationcontinues through forelimb insertion andposteroventrally across the flank where itfades; temporal region dorsally black andlighter gray to tan ventrally, divided alongdiagonal line from anterior end of supratym-panic fold, across center of tympanum, to thepoint of forelimb insertion; lips black withvertically transverse pale cream labial bars,bordered anteriorly by paler brown loweredge of canthus and subocular regions; trans-verse, dark brown bars on limbs similar incolor to dorsum, coalescing into large spots onelbows and knees; three bars on forearms,four on thigh, three on tibia, two on tarsus;mottled flanks, blending into the darker



border (above) and lighter border (below)of dominant color pattern; dorsal surfacesof manus, pes, and digits dark brown withpaler blotches above each joint; dorsal sur-faces of inner two toes slightly paler brownthan outer.

Throat marbled dark brown and dark graywith distinct white flecks; periphery of throatand lower lips darker; chest pale brown withblack wrapping around anteroventral edges ofupper arms; paler gray ventral colorationlimited to thin strip on ventral surfaces offorelimbs; preaxial ventral strips of upper- andforearms cream; venter pale gray with distinct,dark gray spots that decrease in size poster-iorly until blending with uniformly gray groin;ventral surfaces of thigh medium reddishbrown, gray ventral surface of tibia and tarsusuniform brown; palmar and plantar surfaces ofhands and feet dark brown with gray sub-articular tubercles; iris pale gray above anddark gray below pupil.

Coloration of holotype in life.—(differencesfrom preserved specimens observed in digitalphotographs and field notes of CDS andCWL) Dorsum and head dark brown, lackingdorsal color pattern save for dark creaminterorbital bar, slightly paler snout, andscattered white flecks; dorsal surfaces of limbsgolden brown with black transverse bars; headwith golden flecks along edge of canthusrostralis, along vertical labial bars, at tip ofsnout, in subocular region, and along diagonalstripe across black tympanum; dorsal surfacesof digits of manus and pes black with goldenbrown blotches and cross bars at joints; lateralseries of blotches across flanks black; throatand sternal region purplish brown, withdistinct, pale cream spots on venter (fadingposteriorly), groin uniform purplish gray;ventral surfaces of fore- and hind limbsmedium gray with darker peripheries; irisbright gold above and darker golden brownbelow pupil.

Measurements of holotype (mm).—SVL32.9; head length 12.2; head width 14.0; snoutlength 5.2; internarial distance 3.4; interorbitaldistance 3.0; eye diameter 4.1; eye–naresdistance 3.3; tympanic annulus diameter 3.2;eye–tympanum distance 0.5; upper armlength 6.1; forearm length 7.3; femur length17.9; tibia length 18.3; tarsus length 10.0;

manus length 9.3; pes length 18.4; Finger Ilength 6.2; Finger III length 9.3; Toe IVlength 17.2; Finger III disc width 0.9; Toe IVdisc width 1.1; Finger III penultimate pha-lange width 0.7; Toe IV penultimate phalangewidth 0.7.

Variation.—We observed three dorsal colorpatterns in our sample of 15 male specimens.The holotype, eight paratopotypes (PNM9240–41, 9242, 9243; KU 300207, 300210–12), and two paratypes (KU 300204–05) havea dorsum mottled with shades of dark brown,grayish tan, and light hues of brownish green(Fig. 4A). Two paratopotypes (KU 300206,300213) are distinctively colored with a rusty-red and orange dorsum with small patches ofblack and a sharp stratification between thered dorsum and dark flanks (Fig. 4B). Theventral half of the flanks are marbled brown,tan, and brownish green. Rictal tubercleclusters vary from orange to red, and limbsand dorsals surfaces of the hands and feethave irregular patches of red and orange.There is a distinct orange patch on the heels.Two other paratopotypes (KU 300208–09)have two light cream, dorsolateral stripes,one on each side of the body, running fromthe posterior portion of eye to the hind-limbinsertion. The stripes are widest anteriorly andnarrow posteriorly, and the dorsal surfacesbetween the stripes bears a dark brown andblack mottled pattern (darker than that of thefirst color pattern).

The throat, chest, and belly exhibit twodifferent patterns. The first pattern (PNM9240–41, 9243; KU 300204, 300206–08) ismoderately to darkly mottled with tan, gray,dark brown or black patches irregularly in-terspersed. Posterolateral portions of thethroat have two darker brown patches thatsometimes extend well onto the chest andanterior and ventral portions of the upperarm. The chest and belly are tan with darkerbrown markings ranging from small spots toirregular, medium-sized patches. The secondpattern is observed on PNM 9239, 9242 andKU 300205, 300209–13. The throat, chest,and stomach are homogeneously tan, and onlyon the posterior and lateral portions of thethroat are there slightly darker patches of grayto brown of varying distinctiveness. In KU300213, there is a single dark gray patch



observed at the anterior tip of the throat,ventral to the tip of the snout.

Transverse bars on the limbs vary fromdistinct with sharp edges (PNM 9239–40; KU300204, 300208, 300210–13) to less distinct ordifficult to differentiate from the dark brownground color of the limbs (PNM 9241–43; KU300205, 300206–07, 300209). When bars aredistinct from background limb color, they areintermixed with small, black flecks.

All specimens have five transverse dark barson the thigh, four on the tibia, two on thetarsus, and three on the forearms. Palmarsurfaces of the hands and plantar surfaces ofthe feet are dark gray with lighter subarticulartubercles; supernumerary tubercles vary fromlight to dark gray.

Among the type series (15 males includingholotype), SVL 27.7–34.3 (31.8 6 2.0), headlength 30.9–39.6% (34.6 6 0.02) of SVL, headlength 73.0–91.8% (81.8 6 0.05) of headwidth, eye diameter 70.2–106% (80.6 6 0.1)of snout length, 1.0–1.6 (1.3 6 0.15) 3 eye–nares distance, tympanic annulus diameter0.6–0.8 (0.7 6 0.06) 3 eye diameter, manuslength 47.6–54.5% (51.26 0.02) of pes length,pes length 96.8–105% (101 6 0.02) of tibialength, tibia length 54.5–61.5% (57.4 6 0.02)of SVL.

Advertisement call.—At the type locality,visited during the ‘‘dry’’ season, November–May, calling activity was heard only during,and shortly following, rainfall. Wind andadvertisement calls of congeners did notdisrupt the activity of calling males. Callingactivity began shortly after sunset (1830–1930 h) and lasted 3–5 h. Frogs called in-termittently from either atop limestone out-crops or from within crevices in the rock.Males adjacent to other males called morefrequently. Single males were heard makingintermittent sound that sometimes resembledthe advertisement call and at other timesmerely consisted of a simple tone. Periods ofcalling activity were separated by 3–10 min ofsilence before males would begin calling again.

The following description is based on callsof two collected males (PNM 9239; KU300211) and three males that were notcollected. All advertisement calls were re-corded at the same site and at an ambienttemperature of 25.0 C, with cloacal tempera-

tures measured following recordings for thetwo collected 5 26.2 and 26.0 C, respectively.Because each of the five recordings was madeat the same locality and in almost identicalabiotic conditions, we included all of them inthe following analysis. Variation in measure-ments of acoustic characters is given inTable 2.

Males produce a unique advertisement callthat sounds like a series of sharp whistles (thefirst held longer, sweeping in frequency,followed by a rapid set of short whistles) tothe human ear. The first note is a slightfrequency modulated note terminating witha longer tonal signal, followed by shortamplitude modulated pulses (Fig. 5). Duringrecording segments, males called intermit-tently in call groups. Each group consists of 4–12 (7.3 6 2.5; n 5 17) calls per group.Duration of call groups are 6.7–34.3 (17.8 67.2; n 5 17) s. Call groups are separated byintervals of silence, or intergroup intervals, of7.3–66.3 (24.9 6 16.4; n 5 12) s. Callinggroup rate [(total number of call groups 2 1)/time from beginning of first group to begin-ning of last] ranges from 0.012–0.029 (0.02260.008; n 5 4) groups/s.

Within call groups, calling rate [(totalnumber of calls 2 1)/time from beginning offirst call to beginning of last] is 0.20–0.49 (0.386 0.09; n 5 17) calls/s, with calls deliveredmore rapidly toward the end of the call group(intervals between calls decline through callgroups). With two exceptions, final intervalsbetween calls are shorter than the first, withthe interval length difference [first inter-callinterval length–last inter-call interval length]being 0.71–6.9 (3.1 6 2.3; n 5 15) s. Finalintervals between calls are longer than thefirst intervals between calls during the thirdcall groups in KU 300211 (0.5 s difference)and in the second of three non-voucheredrecordings (0.6 s difference). Call intensity isrelatively homogeneous across calls in eachgroup, with each call throughout call groupsaccounting for 88–100% (96.5 6 0.03; n 5 95)of maximum amplitude.

All calls contain an initial note with slightfrequency modulation, followed by a sequenceof rapid amplitude modulated notes. Thereare 0–6 notes following the elongate initialnote, with an increase in the number of notes



per call across call groups. Call groups usuallybegin with a call containing only one note andconclude with longer calls of between threeand six notes. The number of final call notes is2–6 (3.8 6 1.3; n 5 17). Note-repetition rate[(total number of notes per call 2 1)/timefrom beginning of first note to beginning oflast], (excluding the initial, frequency modu-lated note) decreases in the latter calls of eachcall group. In KU 300211, call groups beganwith initial calls with note-repetition rates of8.0–9.0 (8.5 6 0.5, n 5 6) notes/s, and callgroups end with final calls that have note-repetition rates of 7.8–9.0 (8.3 6 0.4, n 5 7)pulses/s. Similar rates also are observed in thesecond non-vouchered recording, with initialnote-repetition rates of 9.3 (9.3 6 0.0, n 5 3)notes/s, and final calls have note-repetitionrates of 8.3–9.3 (8.96 0.4, n 5 5) notes/s. Theinitial frequency modulated note does not varysignificantly in duration between the first andlast call of each call group; this tonal syllablelasts 0.07–0.18 (0.14 6 0.02, n 5 34) s.

The calls of P. paengi have distinct spectralcomponents (Fig. 5A–D; Table 2), with anemphasized (5 dominant) fundamental fre-quency and multiple harmonics (multiples ofthe fundamental). The FFT (power spectrum)demonstrates that the majority of energy ineach call is found in the fundamental fre-quency. There were two detectable harmonicsin all recordings, except for the holotype, inwhich there were three detectable harmonics.

Distribution.—Platymantis paengi is knownonly from 180–300 m above sea level on themountain known locally as ‘‘Mt. Lihidan’’ (11u249 52.740 N, 122u 069 16.740 E; WGS-84),Municipality of Pandan, Antique Province,NW Peninsula of Panay Island, the Philip-pines (Fig. 1).

Etymology.—We take great pleasure innaming the new species for our friend andcolleague Rafe M. Brown, in recognition of his15 yr of contribution to herpetology, his workon the forest frogs of the genus Platymantis,and for his endless support toward interna-tional collaboration. The genitive masculinename ‘‘Paeng’’ is a Tagalog nickname forRafael. Suggested common name 5 PanayLimestone Frog.

Ecology and natural history.—Platymantispaengi is currently known from 180–300 m


TABLE2.—Su

mmaryof

advertisem

entcallcharacters

forthetworecordings

withvouche

rspecim

ensandthreerecordings

whe

revouche

rspecim

enswereno

tcolle

cted

.Whe

napprop

riate,

samplesize

isprovided

andarang

eisshow

nfollo

wed

bymean6

standard

deviation.

The

firstno

n-vouche

redrecordingconsistedof

only

onecallgrou

pand

thereforecann

otbe

evaluatedforseveralcategories

inthistable.

PNM

9239

KU300211

First

non-vouche

redrecording

Second

non-vouche

redrecording

Third

non-vouche

redrecording

Callspe

rgrou

p(8.0

64.2),n5

2(7.1

62.5),n5

7(7.0

60.0),n5

1(7.0

63.3),n5

5(8.0

60.0),n5

2Intergroup

interval

28.9

s7.3–

28.0

s,(18.86

7.3),n5

6Not

applicable

11.9–4

5.6s,

(22.66

15.5),n5

466

.3s

Callin

ggrou

prate

0.0192

grou

ps/s

0.0294

grou

ps/s

Not

applicable

0.02

58grou

ps/s

0.0122

grou

ps/s

With

in-group

calling

rate

0.43–0

.45calls/s,

(0.446

0.01

),n5

20.31–0

.49calls/s,

(18.86

7.3),n5

60.45

,n5

10.20–0

.43calls/s,

(0.316

0.08

),n5

50.28–0

.49calls/s,

(0.386

0.15),n5

2Intercallinterval

differen

ce0.8–2.1s,(1.5

61.0),

n5

20.7–

5.9s,(2.4

62.4),

n5

61.5s,n5

12.1–6.3s,(4.7

61.9),

n5

42.3–

6.9s,(4.6

63.2),

n5

2Callintensity

88–1

00%

maxim

umam

plitud

e,(94.66

3.8),

n5

14

93–100

%maxim

umam

plitud

e,(97.66

2.0),

n5

44

96–1

00%

maxim

umam

plitud

e,(98.56

1.5),

n5

6

91–1

00%

maxim

umam

plitud

e,(95.16

2.7),

n5

24

94–100

%maxim

umam

plitud

e,(96.36

1.8),

n5

7Final

callpu

lsenu

mbe

r2–

5pu

lses,(3.5

62.1),

n5

22–

5pu

lses,(3.7

61.0),

n5

76,

n5

12–

5pu

lses,(3.4

61.3),

n5

53–

5pu

lses,(4.0

61.4),

n5

2FFTpower

spectrum

peaks

3.6kH

z3.6kH

z3.6kH

z3.0kH

z3.0kH

zHarmon

icpe

aks

6.1,

9.0,

and11

.7kH

z6.4and9.4kH

z6.0and8.6kH

z6.0and9.0kH

z5.7and8.9kH

z


above sea level in a small patch of disturbed,secondary-growth forest in karst habitat.Males were observed calling from either thetop of rock outcroppings or from withincrevices in the rock formations.

Calling males were easily disturbed byhuman presence, and we found it difficult toapproach an individual without the frogescaping into limestone crevices. No eggmasses, females, or juveniles were found;other species of Platymantis have directdevelopment (Brown and Alcala, 1982a,b).

Other sympatric anuran species includeLimnonectes visayanus, Platymantis dorsalis,P. corrugatus, P. negrosensis, Polypedatesleucomystax, Kaloula cf. kalingensis, Bufomarinus, and Occidozyga laevis. Amphibiansencountered at nearby lower elevations inmore disturbed habitats included Fejervaryavittigera and Rana erythraea (Ferner et al.,2001; Gaulke et al., 2003).

Conservation.—Platymantis paengi is knownfrom a restricted portion of disturbed habitat inthe NW mountain range of Panay Island. Thespecies was observed between 180–300 mabove sea level; however, we suspect that thespecies may possess a wider elevational rangeand geographical distribution. It is our assump-tion that one of the main limiting factors to thisspecies’ range is distribution of limestoneforest. In the region surveyed, we witnessedwidespread habitat destruction from agricul-tural fields and local farms, and all forestpatches in which we observed the new specieswere regenerated second growth. Currently,we recognize our inability to make a properconservation assessment for this new speciesbecause of a lack of adequate surveys. Wesuggest immediate surveys of karst habitat beconducted across Panay Island in the rainyseason. Such efforts would enable conservationassessments of Platymantis paengi as well asother obligate limestone species.

DISCUSSION

With the new species described here, thereare 27 species of Philippine Platymantis. Thenumber of Philippine Platymantis has in-creased in several bursts, from seven species(Inger, 1954; Brown and Alcala, 1970a,b), to12 species (Brown and Alcala, 1982a), to 25–27 species (Alcala and Brown, 1998, 1999;

Brown et al., 2002; R. Brown, personalcommunication). We expect that this total isstill an underestimate of the actual diversity ofPhilippine Platymantis (Brown et al., 2002; R.Brown, A. Diesmos, and A. Alcala, unpub-lished data). The Philippine radiation ischaracterized by a high degree of documentedcryptic diversity (Alcala et al., 1998; Brown etal., 1997b, 1999), which has only recentlybegun to be resolved because of analyses ofadvertisement-call diversity, microhabitatpreferences, molecular evidence, and mor-phological characters that are currently rec-ognized as important in the delineationbetween species groups and species bound-aries (R. Brown, A. Diesmos, and A. Alcala,unpublished data).

Platymantis paengi is the third Philippinelimestone frog to be described. Others includeP. spelaeus from SW Negros Island and P.insulatus from the Gigantes Islands. Thesespecies appear to have restricted microhabitatpreferences for limestone forest habitat, andso far all occur in the Visayan (Central)Philippine islands. Within these limited geo-graphic ranges, the species are quite commonin the rainy season, but can seem to be quiterare in times of high temperature and aridity(Brown and Alcala, 2000; Brown et al., 2003).

The heavily mottled, dominant morphotypeof Platymantis paengi is similar in appearanceto both P. spelaeus and P. insulatus. With allthree species having unique preferences forrocky, limestone habitat, the similarity in colorpattern may be a result of convergentselection. All three frogs have expandedterminal digital discs; however, P. insulatushas widely expanded discs, a feature that alliesthe species with the P. guentheri Group(sensu Brown et al., 1997a). It is unclear asto which species Platymantis paengi is mostclosely related; a comprehensive molecularphylogenetic analysis will be needed to testcurrent species-group relationships (R.Brown, A. Diesmos, and A. Alcala, unpub-lished data).

The discovery of a new species of limestonefrog within the NW peninsula of Panay Islandhighlights this region as an important compo-nent of the diversity of amphibians andreptiles within the Visayan Islands and thePhilippines. Although surveys at varying



elevations in forested sites are still neededbefore accurate statements on the diversity ofPhilippine Platymantis can be made, wesuggest that simultaneous efforts be focusedon surveys within fragmented and forest edgehabitat throughout the Visayan Islands andthe Philippines. Additional surveys conductedin limestone habitats of Panay are necessary todocument fully the geographic range of thisnew species.

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 C.Custodio and A. Tagtag. We especially thank E. Curio, M.Gaulke, and T. Kuenzel (Philippine Endemic SpeciesConservation Project, PESCP), whose logistical supportand guidance were instrumental throughout visits to NWPanay. We thank the Municipal Mayor of Panay, P.E.Sanchez VI, M. Vengano, E.U. Yengano, and A.D. Rana ofthe Antique Provincial Council and EnvironmentalNatural Resources Office, and B. Offenberger. ArthurOng, L. Averia, V. Yngente, M., Yngente, and N. Antoqueassisted in the field. A special thanks is due to M.Pedigrosa for assistance in obtaining local permission toconduct research in NW Panay. Financial support forfieldwork was provided by the Fulbright Foundation, thePhilippine–American Education Foundation (to CDS),The University of Kansas Natural History Museum andBiodiversity Research Center, and NSF EF-0334952 to L.Trueb and R. Brown. For the loans of specimens(museum abbreviations follow Leviton et al., 1985), wethank 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), D. Cannatella andT. LaDuc (TNHC). Critical reviews of the manuscriptwere provided by R. Brown, L. Trueb, J. Weghorst, E.Lehr, J. Guayasamin and J. Esselstyn. CDS thanks A. C.Alcala and family, G. Alcala, B. Stockwell, J. Maypa, theSUAKCREM staff, A. Ong, L. Averia, A. Diesmos, and M.Diesmos for their support throughout his year in thePhilippines. Finally, we thank R. Brown for the encour-agement and support he has provided to our work in thePhilippines.

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.Accepted: 12 March 2007

.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 banahao.—(6) LUZON ISLAND, QUEZON PROVINCE, Municipality ofTayabas, Baragany Lalo, Mt. Banahao: TNHC 61968–71,PNM 9248–49. Platymantis cagayanensis.—(9) LUZONISLAND, CAGAYAN PROVINCE, ‘‘Tagat Forest Reserve nearSanta Praxedes Town:’’ PNM 7564, 7578, 7496–99, 7506,7608, 7526. Platymantis cornuta.—(2) LUZON ISLAND,KALINGA PROVINCE, Municipality of Balbalan, BarangayBalbalan: CAS 61476 (Holotype); CAS 231498. Platyman-tis corrugatus.—(22) CAMIGUIN ISLAND, CAMIGUIN

PROVINCE, Municipality of Guinsiliban, Barangay Cabuan:KU 300351, 300355; POLILLO ISLAND, QUEZON

PROVINCE, Municipality of Polillo, Barangay Pinaglubayan,62 m elevation, 14u 459 9.30 N, 121u 589 5.520 E: KU300350, 300352–54; NEGROS ISLAND, NEGROS ORIEN-

TAL PROVINCE, Municipality of Valencia, Barangay Bong-bong, Camp Lookout, Cuernos de Negros Mt. Range, Mt.Talinis, 500 m elevation: TNHC 61972–87. Platymantisdorsalis.—(22) NEGROS ISLAND, NEGROS ORIENTAL

PROVINCE, Municipality of Valencia, Barangay Bongbong,Sitio Tagaytay, Mt. Talinis, 1150 m elevation, 9u 159 360N, 123u 129 1960 E: KU 300356–300377. Platymantisguentheri.—(8) BOHOL ISLAND, BOHOL PROVINCE,Municipality of Carmen, Barangay Riverside: TNHC56407; Municipality of Bilar, Barangay Logarito: PNM9250–56. Platymantis hazelae.—(13) NEGROS ISLAND,NEGROS ORIENTAL PROVINCE, Municipality of Valencia,Barangay Bongbong, Sitio Tagaytay, Mt. Talinis, 1150 melevation: KU 300403–15. Platymantis indeprensis.—(8)LUZON ISLAND, QUEZON PROVINCE, Municipality ofTayabas, Barangay Lalo, Mt. Banahao: TNHC 061956–60;PNM 9257–59. Platymantis insulatus.—(12) SOUTHGIGANTE ISLAND, ILOILO PROVINCE, Municipality ofCarles, Barangay Gabi: CAS 117441 (Holotype), 119967–69 (Paratypes); KU 300338–44, 300346; NORTH GI-GANTE ISLAND, ILOILO PROVINCE, Municipality ofCarles, Barangay Granada: KU 300345, 300347–49.Platymantis isarog.—(7) LUZON ISLAND, CAMARINE

SUR PROVINCE, Municipality of Naga City, BarangayPanicuason, Mt. Isarog National Park, Mt. Isarog: TNHC61961–67. Platymantis lawtoni.—(1) TABLAS ISLAND,ROMBLON PROVINCE, Municipality of San Agustin, Mt.



Progreso: CAS 135733. Platymantis levigatus.—(15)SIBUYAN ISLAND, ROMBLON PROVINCE, Municipalityof Magdiwang, Barangay Talaba, Mt. Guiting-GuitingNatural Park, 0 m elevation: KU 300416–30. Platymantisluzonensis.—(16) 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, Municipality ofNaga City, Mt. Isarog: FMNH 251643–44; TNHC 62006–09. Platymantis mimulus.—(12) LUZON ISLAND, LA-

GUNA PROVINCE, Municipality of Los Banos, BarangayBatong Malake, Mt. Makiling: TNHC 54930–31; PNM9260–69. Platymantis montana.—(13) LUZON ISLAND,QUEZON PROVINCE, Municipality of Tayabas, BarangayLalo, Mt. Banahao: TNHC 62149–58; CAS 200998–1000.Platymantis naomiae.—(3) LUZON ISLAND, QUEZON

PROVINCE, Municipality of Tayabas, Baragany Lalo, Mt.Banahao: TNHC 62169–71. Platymantis negrosensis.—(7)NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Cuer-nos de Negros Mountain Range, Mt. Talinis, 750 melevation: KU 300439–45. Platymantis panayensis.—(2)PANAY ISLAND, AKLAN PROVINCE, Municipality ofNabas: CAS 137641–42. Platymantis pseudodorsalis.—(4) LUZON ISLAND, QUEZON PROVINCE, Municipality ofTayabas, Baragany Lalo, Mt. Banahao: KU 207455–57,207459 (Paratypes). Platymantis pygmaeus.—(5) LUZON

ISLAND, CAGAYAN PROVINCE, Municipality of Calveria,Barangay Mabnang, Mabnang Falls: PNM 7523; SIBU-YAN ISLAND, ROMBLON PROVINCE, Municipality ofMagdiwang, Barangay Talaba, Mt. Guiting-Guiting Nat-ural Park, 0 m elevation, 12u 279 40.380 N, 122u 399 51.660E: KU 300431–34. Platymantis rabori.—(3) MINDANAOISLAND, NEW BATAAN PROVINCE, Mt. Puting Bato:CMNH 2305, 2350; DAVAO CITY PROVINCE, Municipalityof toril, Barangay Baracatan, Sitio Upper Baracatan:CMNH 1462. Platymantis sierramadrensis.—(6) LUZONISLAND, AURORA PROVINCE, Municipality of San Luis;Dipiningan branch of the Cobatangan River drainage; 15u409 120 N, 121u 209 480 E: CMNH 5678–79, 5904; ISABELAPROVINCE, Municipality of Palanan, Barangay Didian, SitioNatap Dukan, Northern Sierra Madre National Park, 16u579 55.80N, 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

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

INCE, MtData: CAS 204319–204321. Platymantis taylori.—(4) LUZON ISLAND, ISABELA PROVINCE, Municipality ofPalanan, Barangay Didian, Sitio Natapdukan: CAS207443–207446 (Paratypes).



A NEW SPECIES OF PLATYMANTIS (AMPHIBIA: ANURA…€¦ · A NEW SPECIES OF PLATYMANTIS (AMPHIBIA:...

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