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Aug 14, 2005 - from the Bicol Peninsula of Luzon Island, Philippines. RAFE M. ... significance as a minor center of herpetological diversity and endemism on southern. Luzon. ... and, to some extent, advertisement call (Alcala et al., 1998 ...
COPEIA

2007, No. 2

May 16

Copeia, 2007(2), pp. 251–266

A New Forest Frog of the Genus Platymantis (Amphibia: Anura: Ranidae) from the Bicol Peninsula of Luzon Island, Philippines RAFE M. BROWN

AND JUAN

CARLOS GONZALEZ

We describe a new species of forest frog (genus Platymantis) from Mt. Malinao, Bicol Peninsula, southern Luzon Island, The Philippines. The new species is distinguished from congeners by a moderate body size (28.3–39.1 mm for eight males; 49.8–52.7 in two females), slightly expanded terminal finger and toe disks, a distinct color pattern, a unique microhabitat preference, and various spectral and temporal aspects of the advertisement call. The new species is known only from 950–1160 m above sea level on Mt. Malinao and, as such, accentuates this mountain’s unrecognized conservation significance as a minor center of herpetological diversity and endemism on southern Luzon. We compare the new species to all presumably related (phenotypically similar) species of Platymantis from the Philippines and comment on apparent trends in morphological evolution and habitat preference in Philippine members of the genus Platymantis. Current understanding of Platymantis species diversity throughout this topographically and geographically complex archipelago is underestimated and in need of comprehensive review. Inilalarawan namin ang bagong uri ng palakang gubat (genus Platymantis) mula sa mababa hanggang sa kalagitnaang bulubunduking dipterocarp ng pagbabagong sonang gubat ng Bundok Malinao sa Peninsula ng Bicol sa katimugang pulo ng Luzon, Pilipinas. Ang bagong uri ay may katangiang katamtamang pangangatawan (28.3–39.1 mm para sa walong lalake; 49.8–52.7 para sa dalawang babae), bahagyang malawak na bahaging mabilog at patag sa dulo ng daliri ng kamay at paa. Ito ay may katangi-tanging kulay dibuho, may walang kahambing na mumunting bahay hayop na kanilang higit na nagustuhan, ang iba’t ibang espektrum at pamanahong anyo na kanilang pantawag pansin. Ang bagong uri ay kilala lamang mula sa kalagitnaan ng matayog na lokalidad sa Bundok Malinao, Peninsula ng Bicol at binibigyang diin ang hindi napapansing kahalagahan ng pag-aalaga sa nasabing kabundukan bilang menor na sentro sa pag-aaral ng sari-saring ampibya at reptilya na sa katimugang Luzon lamang matatagpuan. Inihahalintulad namin ang bagong uring ito sa lahat ng inaakala naming may kaugnayan (magkatulad na morpolohiya) sa uri ng Platymantis mula sa Pilipinas at binibigyang puna ang malinaw na takbo ng ebolusyong morpolohiya at ang bahay hayop na higit na nagustuhan sa kaanib na genus sa Pilipinas. Ang kasalukuyang pagkakaintindi sa iba’t ibang uri ng Platymantis sa buong kapuluang may hindi maunawaang topograpiya at geograpiya ay ipinagwawalang bahala. Ito ay nangangailangan ng kabuuang pagbabago.

OUTHEAST Asian forest frogs of the genus Platymantis exhibit one major center of endemism and diversity in the Solomon (Brown, 1952), Bismarck (Zweifel, 1960, 1975; Foufopoulos and Brown, 2004), and Admiralty archipelagos (21 species; Allison, 1996; Allison and Kraus, 2001) and another in the Philippines (26 species; W. Brown et al., 1997a; Alcala and Brown, 1998, 1999). Lesser centers of endemism include intervening islands of Palau (one species; Crombie and Pregill, 1999), New Guinea (six species;

S

#

Zweifel, 1969; Allison, 1996; Gu¨nther, 1999), eastern Indonesia (two to four species; Menzies, 1982a, 1982b; Edgar and Lilley, 1993), and Fiji (two species; Gorham, 1965; Morrison, 2003). In the Philippines, the recognition of three species groups of Platymantis is based on conspicuous characteristics of external morphology (W. Brown et al., 1997a, 1997b, 1999a), ecology, and, to some extent, advertisement call (Alcala et al., 1998; Alcala and Brown, 1999; R. Brown et al., 2003). Most species readily fall into the species

2007 by the American Society of Ichthyologists and Herpetologists

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COPEIA, 2007, NO. 2

groups of W. Brown et al. (1997a), but a few species have strained the boundaries of these assemblages by possessing combinations of characters typical of one or more of the supra-specific groupings. In this paper we describe a distinctive new species that spans the available diagnoses of two species groups. On 11 August 2001, while conducting survey work on the north side of Mt. Malinao, Renato B. Fernandez and RMB collected the first specimens of a remarkably distinct new species of Platymantis at 950 m above sea level (Fig. 1). Later the same year RBF and JCG collected additional specimens on the east side of the mountain, in the municipality of Malinao. The new species combines characteristics of the Platymantis dorsalis group and the Platymantis guentheri group (W. Brown et al., 1997a) by having a terrestrial microhabitat preference (typical of ground frogs of the Platymantis dorsalis species group), slightly expanded terminal disks of fingers and toes, and an amplitude modulated advertisement call (more reminiscent of the tree frogs of the Platymantis guentheri group). Taken together, this and other new species awaiting description (R. Brown, A. Diesmos, and A. Alcala, unpubl. data) suggest that the species groups of W. Brown et al. (1997a) will require eventual revision with the addition of new Philippine taxa to Platymantis. MATERIALS AND METHODS We recorded morphometric data from fluidpreserved specimens in museum collections. Institutional abbreviations follow Leviton et al. (1985), except for the Natural History Museum of the University of the Philippines at Los Ban ˜ os (UPLB-MNH). Sex was determined by gonadal inspection when possible, and measurements were taken (to the nearest 0.1 mm) with digital calipers. In order to minimize inter-observer bias and other sources of potential error (Lee, 1982; Hayek et al., 2001), we used only data scored by RMB. Character definitions follow Zweifel (1960, 1969, 1975) unless otherwise noted. They include: snout–vent length (SVL), head length (HL), eye–narial distance (END), snout length (SNL), interorbital distance at the midpoint of the orbits (IOD), internarial distance (IND), eye diameter (ED), horizontal tympanic annulus diameter (TAD), eye–tympanum distance (ETD), head width at the widest point (HW), upper arm length (UA), forearm length (FAL), tibia length (TBL), femur length from the cloaca to the outer surface of the flexed knee (FL),

Fig. 1. Known distribution of Platymantis diesmosi on Mt. Malinao (inset), Bicol Peninsula, Luzon Island, Philippines. The capital city of Manila is indicated with a star, and the type locality (Mt. Malinao, municipality of Tiwi, Albay Province) is indicated by a black dot. The volcanic peak of Mt. Mayon (A) is located immediately SE of Mt. Malinao, and Mt. Isarog (B) is located to the NW. Scale bar 5 10 km.

BROWN AND GONZALEZ—NEW PHILIPPINE PLATYMANTIS tarsus length (TSL), pes length (PL), manus length from tip of third digit to the base of the outer metacarpal tubercle (ML), fourth toe length (Toe4L), first finger length (Fin1L), third finger length (Fin3L), first finger disk width (Fin1DW), third finger disk width (Fin3DW), fourth toe disk width (Toe4DW), and widths of penultimate phalanges of third finger (PpFin3) and the fourth toe (PpToe4). Other morphological characteristics included color pattern, dermal ornamentation, and size and shape of subdigital tubercles. Advertisement calls were recorded with a Sony TM WM DC6 Professional Walkman with a SennheiserTM ME80 condenser microphone (equipped with K3U power module). Calls were recorded at distances of 2–4 m, and ambient, substrate, and cloacal temperatures were collected immediately after call recording. Calls were digitized using Soundedit (Macromedia, 1995) and analyzed using Canary (Charif et al., 1996) software. Temperature corrections were not necessary because frogs were recorded at nearly identical temperatures. We examined oscillograms (waveforms), audiospectrograms, and results of the Fast Fourier Transformation (FFT; power spectrum) for a series of spectral and temporal call characteristics following Diesmos et al. (2002a), R. Brown et al. (2003), and Foufopoulos and Brown (2004). Calling rate, calling group rate, and pulse repetition rate was calculated according to: ([total number of calls/ groups/pulses – 1]/[time from beginning of first group to beginning of last]). Platymantis diesmosi, new species Mt. Malinao Forest Frog Figures 2–3 Holotype.—PNM 8499 (field no. RMB 3526), adult male, Philippines, Luzon Island, Albay Province, municipality of Tiwi, Barangay Banhaw, Sitio Purok 7, area known locally as ‘‘Tamagong,’’ 950 m above sea level, NW slope of Mt. Malinao, 13u269120N, 123u349240E, 2015 hr, 11 Aug. 2001, Renato B. Fernandez and Rafe M. Brown. Paratypes.—TNHC 62040 (RMB 3524), 62041 (RMB 3525), and 62042 (RMB 3527), 3 males, same data and locality as holotype; UPLB-MNH 16, 21–22, PNM 8501, 4 males; UPLB-MNH 23, PNM 8500, 2 females, Philippines, Luzon Island, Albay Province, municipality of Malinao, Barangay Sugcad, area known locally as ‘‘Sugcad Plaza,’’ 1125–1160 m above sea level, small tributary of the Kumagingking River, SE slope of Mt. Malinao, 13u249380N, 123u399130E, 2000–2100 hr, 13 May 2002, R. B. Fernandez, J. C. Gonzales, and party.

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Diagnosis.—Platymantis diesmosi is unique among species of the genus Platymantis by virtue of (1) moderately large body size (28.3–39.1 mm SVL for males; 49.8–52.7 for females), (2) slightly expanded digital disks on tips of fingers and toes, (3) smooth (or only faintly longitudinally-ridged) skin of the dorsum, (4) unique microhabitat preference, (5) distinct coloration, and (6) numerous temporal and spectral characteristics of the advertisement call (Table 1). Although the new species is clearly diagnosable as a member of the Philippine P. guentheri group (W. Brown et al., 1997a, 1997b), we include comparisons to all the relevant (phenotypically similar and/or geographically proximate) species of Philippine Platymantis. From P. pygmaeus, P. naomiae, and P. mimulus, the new species is distinguished by its much larger body size (males: 28.3–39.1 mm vs. 14.0– 15.6 in P. pygmaeus, 20.3–24.0 in P. naomiae, and 18.0–25.8 in P. mimulus), and microhabitat preference (cliff/ravine edge vs. forest floor, and/or herb-layer vegetation). It also is diagnosed by its amplitude modulated advertisement call (vs. single pulse calls in P. pygmaeus and complex, multi-syllable calls in P. naomiae and P. mimulus). From P. dorsalis, P. cagayanensis, P. corrugatus, P. indeprensus, P. pseudodorsalis, and P. levigatus, P. diesmosi is diagnosed by a tendency towards a larger maximum body size (males: 28.3–39.1 mm vs. 25.5–32.2 in P. dorsalis, 26.0– 30.8 in P. cagayanensis, 28.2–33.5 in P. corrugatus, 22.4–31.5 in P. pseudodorsalis, 28.7–30.0 in P. levigatus, and 26.5–33.4 in P. taylori). It also can be distinguished by its amplitude modulated advertisement call (vs. frequency sweeps in P. dorsalis and P. pseudodorsalis and/or complex, multi-syllable calls in the remaining P. dorsalis group species) and from all these species except P. pseudodorsalis, microhabitat preference (cliff/ ravine edge vs. forest floor and/or shrub layer vegetation). Platymantis diesmosi is distinguished from P. spelaeus by its smaller (males: 28.3–39.1 vs.41.5–46.9 mm SVL) body size, microhabitat preference (cliff edge vs. limestone caves and crevices), and the presence of an amplitude modulated (vs. complex, multi-syllable tonal) advertisement call (R. Brown et al., 2003). From frogs of the P. hazelae group (W. Brown et al., 1997a; Alcala and Brown, 1999: P. hazelae, P. isarog, P. lawtoni, P. montanus, P. panayensis, P. polillensis, P. sierramadrensis, P. subterrestris), P. diesmosi is distinguished by a larger body size (males: 28.3–39.1 mm vs. 21.0–26.5 for males of all P. hazelae group species), distinct and pointed (vs. indistinct and flattened) subdigital tubercles of manus and pes (W. Brown et al., 1997a:fig. 1), an unreduced (vs. reduced) first digit of the

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Fig. 2. Photographs in life of Platymantis diesmosi paratypes exhibiting two color patterns: (A) the marbled black phase (TNHC 62042, male, 36.0 mm SVL; Photo: RMB); and (B) the solid brown phase (UPLB-NHM 22; male, 33.8 mm SVL; Photo: JCG).

Fig. 3. Lateral (A) and dorsal (B) view of the head of the Platymantis diesmosi holotype (PNM 8499). Scale bar 5 5 mm.

manus, slightly expanded (vs. widely expanded) terminal disks of the manus and pes, amplitude modulated (vs. tonal) advertisement call, and by terrestrial (vs. exclusively arboreal) microhabitat preference. Platymanis diesmosi is distinguished from the primarily arboreal species of the Platymantis guentheri group (W. Brown et al., 1997a, 1997b: P. banahao, P. cornutus, P. guentheri, P. insulatus [Fig. 4], P. luzonensis, P. negrosensis, P. rabori) by slight (vs. wide) expansion of the terminal disks of the manus and pes (Fig. 4), and with the exception of the terrestrial P. insulatus, by a terrestrial (vs. arboreal) microhabitat preference. The new species is further distinguished from P. guentheri by possession of an amplitude modulated (pulsed) advertisement call (vs. frequency sweep call) and from P. luzonensis, P. negrosensis, P. rabori by a rapid (vs. slow) pulse rate of the advertisement call (Table 1; W. Brown et al., 1997b; R. Brown et al., 2003). Although body

28.3–39.1 Mt. Malinao

Dorsal dermal tubercles 2 Dorsal longitudinal 2, faint folds/ridges Terminal digital disk Narrow expansion Microhabitat preference Terrestrial Advertisement call Rapidly pulsed

SVL (mm) Range

P. diesmosi (8 m)

Arboreal Slowly pulsed

Arboreal Arboreal Frequency sweep Rapidly pulsed

+ 2, + Wide

2 2

+ 2

26.8–36.2 Luzon PAIC

P. luzonensis (12 m)

Moderate to wide Wide

27.3–39.6 Mt. Banahao

P. banahao (12 m)

24.1–32.9 Mindanao PAIC

P. guentheri (11 m)

Arboreal Slowly pulsed

Wide

+ 2, +

29.1–39.1 Visayan PAIC

P. negrosensis (10 m)

Terrestrial Rapidly pulsed

Wide

2 2

37.1–43.8 Gigante Islands

P. insulatus (4 m)

Arboreal Slowly pulsed

Wide

+ 2, +

27.1–33.2 Mindanao PAIC

P. rabori (8 m)

Non-expanded to narrow Terrestrial Frequency sweep

25.5–32.2 Luzon + Visayan PAICs + 2, +

P. dorsalis (13 m)

TABLE 1. SUMMARY OF THE DISTRIBUTION OF QUALITATIVE DIAGNOSTIC CHARACTERS (+ PRESENT, – ABSENT) IN Platymantis diesmosi AND OTHER PHILIPPINE ‘‘TREE FROGS’’ OF THE GUENTHERI SPECIES GROUP, GENUS Platymantis. Sample size, body size of males, and general geographical distribution (PAIC 5 Pleistocene Aggregate Island Complexes, sensu Brown and Diesmos, 2002) are included for reference, as are data for a representative exemplar of the P. dorsalis species Group (P. dorsalis).

BROWN AND GONZALEZ—NEW PHILIPPINE PLATYMANTIS 255

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Fig. 4. Palmar and plantar view of manus (A) and pes (B) of Platymantis dorsalis (TNHC 21870); (C, D) P. diesmosi holotype PNM 8499; and (E, F) P. banahao (PNM 3640). Scale bars 5 2 mm.

size of P. diesmosi widely overlaps that of most other P. guentheri group species (e.g., most species have SVLs of 25–39 mm; W. Brown et al., 1997b), P. insulatus tends towards a markedly larger maximum body size (37.1–43.8 mm SVL) than P. diesmosi (28.3–39.1). Description of holotype.—A mature, undissected male in excellent condition; habitus moderately robust; head only slightly distinct from neck and wider in dorsal aspect than body, length 43.5% of

SVL; head length 92.6% of head width; snout barely protruding beyond lower jaw, tip bluntly pointed in dorsal aspect (Fig. 3), posteroventrally inclined in profile; eyes protruding slightly beyond silhouette of head in dorsal aspect, well above dorsal surface of head in profile; lips flared and swollen, extending slightly beyond most lateral edge of orbits when viewed in dorsal aspect; interorbital region flat; eye diameter slightly wider than interorbital distance; pupil horizontally elliptical in preservative, horizontally

BROWN AND GONZALEZ—NEW PHILIPPINE PLATYMANTIS elliptical in life; canthus rostralis nearly straight; loreal region strongly concave, with ventral portions sloping ventrolaterally to wide labial shelf; eye diameter 78.6% of snout length, 1.3 times eye–narial distance; nostrils laterally protuberant, oriented posterolaterally; eye–narial distance 3.5 times longer than distance from nostril to tip of snout; internarial region slightly convex; tympanic annulus distinct, its diameter 67.3% of eye diameter; dorsal edge of tympanic annulus sightly concealed by supratympanic fold, the latter extending from dorsal edge of tympanum to supra-axillary region; tongue elongate, subtriangular (widest posteriorly), with deep posterior notch and narrow anterior attachment; choanae situated at anterolateral edge of palate, partially concealed by palatal shelf of the maxillae, round, small, widely separated by a distance five to six times greater than their diameters; dentigerous process of vomer distinct, with six teeth; dentigerous process angled anterolaterally, with closest (posterior) points separated by a distance slightly greater than a single choana, their most distant (anterior) ends separated by a distance equal to three times the diameter of a single choana; openings to vocal sac short slits. Skin of dorsal surfaces of body, head, and limbs finely granular; dermal ornamentation including elongate, very slightly pronounced dorsal ridges, concentrated in suprascapular region; ventral surfaces of trunk, head, throat, and limbs smooth, save for glandular texture concentrated on the posterior 1/4 of trunk and medial 1/3 of thighs; limbs, digits, manus, and pes well developed (Fig. 5C–D), proportionally long; manus length 54.5% of pes length; pes length 98.6% of tibia length; femur 94.6% of tibia length; tibia length 57.8% of snout–vent length; digits of manus slender, long; terminal disks slightly to moderately expanded (Fig. 4C) with pronounced distal, crescent-shaped circummarginal folds; dorsal surfaces of terminal phalanges with cutaneous supra-articular flaps between ultimate and penultimate phalanges; relative lengths of fingers I 5 II , IV , III; subarticular tubercles prominent, convex to pointed on their ventral surfaces, velvety in texture; one subarticular tubercle on digits I–II, two tubercles under digits III–IV; supernumerary tubercles present at the base of digits II–IV; supernumerary tubercles of digit IV flat, indistinct; of digits II–III distinct, round, prominent; thenar (inner metacarpal) tubercle extremely large, elongate, oval (longer axis parallel to digital axis); ventromedial edge raised, sharp, protruding beyond inner (medial) edge of digit I; thenar tubercle much longer than rounded

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palmar tubercle; palmar (outer metacarpal) tubercle nearly divided into medial and outer metacarpal tubercles; medial tubercle large, subcircular, slightly convex; outer metacarpal tubercle elongate, less than half the size of medial metacarpal tubercle; nuptial pads absent, forearms slender, musculature not hypertrophied. Tarsus smooth, lacking folds, flaps, or tubercles; terminal disks of toes moderately expanded, with circummarginal grooves and supra-articular cutaneous flaps above ultimate–penultimate phalangeal articulation; plantar surfaces of pes smooth, with moderately-developed, prominently rounded subarticular tubercles (Fig. 4D); relative lengths of toes I , II , V , III, IV; outer metatarsal tubercle small, round, pointed; inner metatarsal tubercle prominent, with sharp plantar edge, as long as distance from base of toe I to first subarticular tubercle; toes barely webbed basally (not extending to the level of the proximal subarticular tubercles). Cloacal region glandular, lacking tubercles or supracloacal flaps. Coloration in preservative.—Dorsal ground color on head and body nearly homogeneous dark brown with distinct pale gray patches; dorsal surfaces of limbs medium brown with transverse dark brown bars; faint, slightly paler interorbital bar bordered posteriorly by dark brown and anteriorly by slightly paler snout; wide black bar extending from nares, through canthal, preocular, postocular, and tympanic regions, wrapping ventrally around posterior edge of tympanic annulus (where expanded so as to cover anteroventral edge of tympanum), continuing through forelimb insertion, and posteroventrally across the flank where it fades; temporal region black with slightly paler ventral edge, giving impression of diagonal, oblong, white oval, with longer axis extending posteroventrally across tympanum; lips black with vertically transverse pale cream labial bars, bordered anteriorly by paler brown lower edge of canthus and subocular regions; transverse dark brown bars on limbs similar in hue to dorsum, and coalesced into large spots on elbows and knees; three bars on forearms, four on thigh, three on tibia, two on tarsus; horizontal series of fused black spots on flanks (largest, darkest, and most prominent in the immediate postaxial region) extending across flank at the stratified border of the darker (above) and lighter (below) color pattern; dorsal surfaces of manus, pes, and digits dark brown with paler blotches above each joint; dorsal surfaces of inner two toes slightly paler shade of brown.

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Fig. 5. The advertisement call of Platymantis diesmosi (paratype TNHC 64042) recorded at ambient temperature of 22.2 C (cloacal temp. 24.0 C) on 11 August 2001: power spectrum (A: Fast Fourier Transformation; relative amplitude vs. frequency in kHz) and an expanded waveform (B: relative amplitude vs. time in ms) of a single call. The sonogram (C: audiospectrogram: frequency in kHz vs. time in s) and an oscillogram (D: relative amplitude vs. time in s) of the final four calls from a single call group. Background signals visible at 1.5, 3–4, and 5 kHz (in C) are insect vocalizations; the four-pulsed call of Platymantis luzonensis (2.1 kHz) overlaps the first P. diesmosi call in this figure. Expanded inset in waveform (B) shows subpulse structure over 100 ms.

Throat marbled dark brown and dark gray with distinct white flecks; periphery of throat and lower lips distinctly darker; chest pale brown with black wrapping around anterior edges of upper arms; paler ventral coloration limited to thin strip on ventral surfaces of forelimbs; central ventral strips of upper- and forearms cream, with darker periphery; venter pale gray

with distinct dark gray spots that decrease in size posteriorly until blending with uniformly gray groin; ventral surfaces of thigh medium reddish brown, gray ventral surface of shank and tarsus homogeneous brown; palmar and plantar surfaces of hands and feet dark brown with gray subarticular tubercles; iris pale gray above and dark gray below pupil.

BROWN AND GONZALEZ—NEW PHILIPPINE PLATYMANTIS Coloration in life.—(from color transparencies and field notes of RMB; Fig. 2) Dorsum and head dark brown, lacking dorsal color pattern save for dark cream interorbital bar, slightly paler snout, and scattered white flecks; dorsal surfaces of limbs golden brown with black transverse bars; head with golden flecks along edge of canthus rostralis, along vertical labial bars, at tip of snout and in subocular region, and along diagonal stripe across black tympanum; canthal region, pre- and postocular regions, and supra- and posttympanic regions almost black; dorsal surfaces of digits of manus and pes black with golden brown blotches and cross bars at joints; inner two digits of both manus and pes solid dark gray, lacking golden brown pigment; lateral series of blotches across flanks black; throat and sternal region purplish brown, with distinct pale cream spots on venter (fading posteriorly), groin uniform purplish gray; ventral surfaces of fore- and hindlimbs medium gray with darker peripheries; palmar and plantar surfaces of hands and feet purplish gray with pale gray tubercles; iris bright gold above and darker golden brown below pupil.

TABLE 2. MORPHOMETRIC VARIATION IN THE TYPE SERIES Platymantis diesmosi. Table entries include mean (6 1 SD), above, and range, below.

OF

Character

Males (8)

Females (2)

Holotype (m)

SVL

35.6 6 3.7 28.3–39.1

51.3 6 2.0 49.8, 52.7

38.2

HL

15.6 6 1.7 12.7–17.7

20.5 6 0.02 20.5, 20.6

16.6

HW

14.5 6 1.3 12.2–16.1

21.0 6 0.2 20.9, 21.2

15.2

SL

7.7 6 0.8 5.9–8.2 4.0 6 0.5 3.2–4.7

9.3 6 0.5 9.0, 9.7 5.0 6 0.2 4.9, 5.2

7.0

IOD

3.9 6 0.5 3.3–4.7

5.1 6 0.1 5.0, 5.2

4.0

ED

5.0 6 0.8 4.0–5.7

6.7 6 0.1 6.6, 6.8

5.5

END

4.7 6 0.5 3.7–5.3

6.7 6 0.7 6.2, 7.1

4.2

TAD

3.2 6 0.6 1.9–3.8 1.6 6 0.2 1.1–1.8

3.5 6 0.4 3.2, 3.8 2.3 6 0.1 2.3, 2.4

3.7

UA

7.4 6 0.8 6.2–8.9

9.6 6 0.2 9.4, 9.7

7.7

FAL

9.2 6 0.9 7.2–10.1

14.0 6 0.03 13.9, 14.1

9.4

FL

20.4 6 1.3 17.6–21.2

29.6 6 0.7 29.1, 30.1

20.9

TBL

21.5 6 1.4 18.9–22.9 12.1 6 0.6 11.0–12.8

31.6 6 0.2 31.4, 31.7 17.3 6 0.3 17.2, 17.5

22.1

ML

10.5 6 1.1 8.9–11.9

15.3 6 0.4 15.0, 15.6

11.9

PL

21.4 6 1.7 18.2–22.7

29.2 6 1.3 28.2, 30.1

21.8

Fin1L

5.8 6 0.5 5.0–7.0

9.1 6 0.6 8.7, 9.5

5.7

Fin3L

7.6 6 0.8 6.4–8.9 15.7 6 1.4 13.7–18.0

11.0 6 0.3 10.2, 10.8 23.9 6 0.4 23.6, 24.2

9.0

Fin3DW

1.1 6 0.1 0.9–1.3

1.1 6 0.1 0.9, 1.3

1.2

Toe4DW

1.1 6 0.1 0.9–1.3

1.7 6 0.3 1.6, 1.8

1.2

PpFin3

0.8 6 0.2 0.5–1.1

1.1 6 0.04 1.0, 1.1

0.8

PpToe4

0.7 6 0.07 0.6–0.9

1.1 6 0.2 1.0–1.2

0.6

IND

ETD

Variation.—Morphometric variation in the type series is presented in Table 2. There are three basic dorsal color patterns in our sample of ten specimens. In the holotype and five paratypes (TNHC 62041; UPLB-MNH 16, PNM 8501, UPLB-MNH 21–22) the dorsum is brown with minute white flecks and/or irregular darker brown blotches. In two paratypes (TNHC 62040, 62042), the dorsum is nearly black, interrupted with an indistinct marbling of brown on the dorsolateral surfaces of body, head, and limbs. In this second pattern, highlights of gold are particularly intense around the edges of the palpebra, on the snout, and in temporal and loreal regions. Additionally, gold pigment coalesces into transverse bars of the limbs and digits. Finally, in one female paratype (UPLB-MNH 23), the dorsum is khaki tan with thick, indistinct yellowish straight dorsolateral stripes, and a distinct narrow white vertebral stripe. The throat varies from nearly homogeneous dark brown (TNHC 62042) to pale brown with white blotches (TNHC 62041) to cream with slight brownish-yellow marbling (UPLB-MNH 16 [male], and 23 [female]). The chest and belly varies from dark with minute white spotting (TNHC 62041) to white with dark brown blotches (TNHC 62042), to pale cream (UPLBMNH 16). In life, two specimens (TNHC 62041– 42) had faint bluish coloration in the immediate axillary region. The cloacal region has an inverted, crescent-shaped white mark in some

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TSL

Toe4L

4.3

1.7

12.5

15.1

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specimens (TNHC 62041–42) or has indistinct white spots in others (TNHC 62040, PNM 8499). The transverse bars of limbs vary from distinct (TNHC 62042) to less evident and fragmented (TNHC 62040, PNM 8499). They also vary from wavy (TNHC 62042, UPLB-MNH 16, 20) to nearly straight (TNHC 62041, UPLB-MNH 23). All specimens have four transverse dark bars on the thigh and three on the shank except TNHC 62042, which has six on the shank. All specimens have three transverse bars (albeit of varying distinctiveness) on the forearm. Owing to variation of the density of melanophores on the palms, supernumary tubercles vary from clearly distinct (TNHC 62040, 62042, PNM 8500, UPLBMNH 23) to obscure and less perceptible to the unaided eye (TNHC 62041, UPLB-MNH 16, 20, 22). Advertisement call.—At the type locality, calling activity was not heard during heavy rain or high levels of ambient noise from heavily running rivers or wind. Only on relatively ‘‘quiet’’ nights following rain did calling activity occur. On these nights, calling activity began approximately one hour after sunset (1900 hr) and lasted for 2– 4 hours. Frogs called intermittently from edges of cliffs in loose, chorus-like organization (when 2–6 individuals were audible to a single observer), interspersed with 5–15 minutes of silence before adjacent individuals on the same cliff stimulated one another to call again. Males of Platymantis diesmosi produce a unique advertisement call in the form of a rapid amplitude modulated group of notes (Fig. 5), sounding to the human ear like a series of harsh, nontonal, accelerating clicks, knocks, or crunches: ‘‘Er . . . er . . . er–er–ererer . . . ’’ The following description is based on calls of two males (PNM 8499 and TNHC 62042) recorded at ambient temperatures of 22.2 and 22.4 C, respectively (cloacal temperatures following recording 5 23.7 and 24.0 C, respectively). During recording segments of 15–20 m, P. diesmosi called in multiple bouts (5call groups), with 3–6 calls per call group (x¯ 5 4.2 6 0.6 SD; n 5 6 for PNM 8499; x¯ 5 4.4 6 0.8; n 5 7 for TNHC 62042). Duration of call groups were 3.5– 9.8 (x¯ 5 6.2 6 1.8 SD; n 5 7) s in PNM 8499 and 3.6–9.3 (x¯ 5 6.2 6 2.2 SD; n 5 6) s in TNHC 62042. PNM 8499 called in groups separated by intervals of silence of 104–216 (x¯ 5 141 6 42.0 SD; n 5 6) s; TNHC 62042 called in groups separated by intervals of 92–132 s (x¯ 5 113 6 16.5 SD; n 5 5). Calling group rate was 0.0053 groups/s for PNM 8499 and 0.0061 groups/s for TNHC 62042.

Within call groups, calling rate was 0.49–0.57 (x¯ 5 0.53 6 0.03 SD; n 5 7) calls/s for PNM 8499 and 0.53–0.58 (x¯ 5 0.55 6 0.02 SD; n 5 7) calls/s in TNHC 62042. Within call groups, calls were delivered more rapidly towards the end of the call group (intercall intervals declined through call groups). Within call groups, final intercall intervals were always shorter than the first, with the difference being 0.11–0.36 (x¯ 5 0.22 6 0.12 SD; n 5 6) s in PNM 8499 and 0.15–0.52 (x¯ 5 0.34 6 0.14 SD; n 5 5) s in TNHC 62042. Throughout these same call groups, amplitude increases with each call (last calls loudest). Relative call intensity (dB) of each call throughout call groups increased by 17–31% (x¯ 5 24.2 6 7.6 SD; n 5 6) of maximum amplitude in PNM 8499 and by 14–26% (x¯ 5 21.2 6 6.1 SD; n 5 5) of maximum amplitude in TNHC 62042. Calls contained 2–13 dull, low frequency, nontonal notes (Fig. 5B) per call. There was a marked increase in number of notes per call across call groups. Call groups usually began with short calls of only a few notes and ended with drawn out calls containing 8–13 notes. Mean note number for the first call per group was 2–6 (x¯ 5 4.5 6 2.4 SD, n 5 7) in PNM 8499 and 2–4 (x¯ 5 2.7 6 0.5 SD, n 5 6) in TNHC 62042. These same call groups terminated with 8–13 (x¯ 5 11.3 6 2.1 SD, n 5 7) notes per final call in PNM 8499 and 9–13 (x¯ 5 11.1 6 1.8 SD, n 5 6) notes per call in TNHC 62042. Note repetition rate also increases in the latter calls of each call group. In PNM 8499, call groups began with initial calls that had note repetition rates of 5.1–11.5 (x¯ 5 7.1 6 2.2 SD, n 5 7) notes/s and call groups ended with final calls that had note repetition rates of 9.4– 13.3 (x¯ 5 12.1 6 1.3 SD, n 5 7) pulses/s. TNHC 62042 initially called with note repetition rates of 5.0–6.1 (x¯ 5 5.5 6 0.45 SD, n 5 6) note/s, and final calls had note repetition rates of 9.4–13.1 (x¯ 5 11.2 6 0.53 SD, n 5 6) pulses/s. Each note was composed of 3 or 4 amplitude-modulated pulses (Fig. 5B: inset), and from initial pulses to maximum pulse amplitude there was an approximately 50–60% increase in relative percent of maximum amplitude (Fig. 5D). The calls of P. diesmosi have distinct spectral components (Fig. 5A, 5C), including an emphasized fundamental frequency with multiple harmonics. The FFT (power spectrum) demonstrates that the majority of energy in each call is packaged in the fundamental (5dominant) frequency, which peaks between 1.2 (PNM 8499) and 1.4 (TNHC 62042) kHz. Two detectable harmonics (multiples of the fundamental) peaked at 2.4 and 4.8 in PNM 8499, and at 2.8 and 5.5 kHz in TNHC 62042.

BROWN AND GONZALEZ—NEW PHILIPPINE PLATYMANTIS Ecology and natural history.—Platymantis diesmosi is currently known only from mid-elevational transitional forest between lower- and mid-montane dipterocarp forest communities as defined by Whitmore (1984). At the type locality the calling males of the new species exhibited a microhabitat preference unique among Philippine Platymantis. All calling males observed in the municipality of Tiwi that were heard over an eight-day period vocalized solely from the edge of steep cliffs along deep arroyo-like creek ravines. At Tiwi, calling frogs were observed at the upper edges of a few cliffs, 15–35 m above the level of the water below, and were situated only on dry patches of soil directly underneath the overhanging edge of the cliffs’ upper lip. Calling males faced out into the forest, away from the cliff edge, and calls could be heard 50–75 m away in the forest below. In the municipality of Malinao, specimens also were only collected 30– 40 m away from water, at the upper edge of a slope. Despite extensive searches below and above cliff ledges on the plateau above, individuals were only encountered along the narrow strip of exposed soil at the edge of the overhanging and collapsing cliff edge. At the municipality of Malinao, the collection site was predominantly lower montane rainforest. Dominant trees were Podocarpaceae (Podocarpus, Dacycarpus), Lauraceae, some Moraceae, Fabaceae (Lithocarpus) and Melastomaceae (Medenilla), some stilt-rooted Pandanaceae (Pandanus), many epiphytic shrubs, ferns, and orchids with moderate to thick moss cover on branches and trunks. During surveys on Mt. Malinao, biologists spent several weeks at the lower edge of the forest (approx. 700–800 m) and at higher elevations between the type locality and the mountain’s peak (1550 m). Platymantis diesmosi was only encountered between 900 and 1050 m above sea level. Other anuran species detected sympatrically include Limnonectes woodworthi, L. macrocephalus, Platymantis dorsalis, P. corrugatus, P. cf. indeprensus, P. cf. isarog, P. luzonensis, Rhacophorus bimaculatus, Polypedates leucomystax, Philautus surdus, Kaloula cf. kokacii, Occidozyga laevis, Rana (‘‘Chalcorana’’) luzonensis, and Rana (‘‘Pulchrana’’) similis. Amphibians encountered at nearby lower elevations in more disturbed habitats included Bufo marinus, Rana (‘‘Fejervarya’’) vittigera, Kaloula picta, and K. c. conjuncta. Distribution.—The new species is known only from the municipalities of Tiwi and Malinao, Mt. Malinao, Albay Province, Bicol Peninsula, Luzon Island, The Philippines (Fig. 1).

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Conservation.—The new species is presently known only from a narrow elevational band around Mt. Malinao between the elevations of 900 and 1050 m above sea level. Because this elevational stratification corresponds to the transition zone between lower- and mid-montane dipterocarp forests (Whitmore, 1984), we assume that the new species may be limited to this single habitat type. Near the type locality, we witnessed extensive slash-and-burn shifting agriculture and selective logging throughout mid-montane elevations on Mt. Malinao. In 2001, the type locality was barely 100 m above some of the most severe disturbances on the eastern face of Mt. Malinao, and we fear that continued degradation of the forest edge at this site will soon extend upwards to the type locality. Because this area also supplies the majority of fresh water to the municipality of Tiwi, continued and unchecked exploitation of this fragile transition zone habitat will threaten not only the continued existence of Platymantis diesmosi, but also the future availability of fresh water for humans in the Tiwi watershed. Conservation initiatives aimed at protecting habitat of potentially threatened (by virtue of a presumed restricted geographical distribution) populations of P. diesmosi could also guarantee the future of clean water for the human populations surrounding the southern and eastern foothills of Mt. Malinao. Etymology.—We take great pleasure in naming the new species for our friend and colleague Arvin C. Diesmos. Arvin has discovered ten (W. Brown et al., 1997a; Alcala et al., 1998) of the recently described species of Philippine Platymantis (W. Brown et al., 1997b, 1999a, 1999b). He also has identified at least as many currently undescribed species as part of his unceasing attention to the study of Philippine amphibian diversity, conservation, and ecology. DISCUSSION The new species described in this paper brings the number of known species of Platymantis from the Philippines to 27. The cumulative number of Philippine Platymantis has risen sharply over the past decade, from seven species (Inger, 1954; W. Brown and Alcala, 1970b), to 12 species (Allison, 1996; W. Brown, 1997; Inger, 1999), to 25–26 species (Alcala and Brown, 1998; W. Brown et al., 1999a; R. Brown et al., 2002a) and now reapproximates the original estimates of E. H. Taylor (1920, 1922a, 1922b). This comes as no surprise; like Taylor (1923, 1925) we are certain that the total species diversity in the Philippine radiation is still substantially underestimated.

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The potential for cryptic speciation in this group appears high (W. Brown et al., 1997c, 1999a; Alcala et al., 1998). It is only now that we have data on advertisement calls, microhabitat preferences, coloration in life, and subtle differences in morphology that we are beginning to recognize the importance of previously underappreciated characters for delineation of species boundaries in Philippine Platymantis (R. Brown, A. Diesmos, and A. Alcala, unpubl. data). Platymantis diesmosi exemplifies the complexity of morphological and behavioral adaptations that we have only recently begun to appreciate in Philippine Platymantis. The new species is a terrestrial frog, presumably related to an almost exclusively arboreal group of frogs, and yet it possesses a combination of attributes unique among all Philippine species of the genus Platymantis. The only other terrestrial frog of the P. guentheri tree frog species group (sensu W. Brown et al., 1997a) is P. insulatus, which is endemic to the tiny land-bridge islands of the Gigante group (W. Brown and Alcala, 1970a; R. Brown and Alcala, 2000; Ferner et al., 2001) of the Visayan Pleistocene Aggregate Island Complex (W. Brown and Alcala, 1970a; R. Brown and Diesmos, 2002). Among species of the Platymantis guentheri species group, P. insulatus is uniquely adapted to a terrestrial limestone habitat (R. Brown and Alcala, 2000; R. Brown et al., 2003). Platymantis diesmosi, in contrast, is a forest species endemic to the Bicol Peninsula, where it inhabits moist lower montane forests and has a highly specific microhabitat preference on the edges of a few sheer cliffs on drainage ridges of Mt. Malinao. We suspect that the phenotypic similarity of these two species is an artifact of convergent selection and that the two will eventually be identified as unrelated ecological specialists. Testing of this hypothesis will require an independent (i.e., molecular) phylogenetic estimate for Philippine species of Platymantis and a comprehensive survey of morphological and acoustic variation in Philippine populations (R. Brown, A. Diesmos, and A. Alcala, unpubl. data). The presence of a new species of terrestrial ‘‘tree frog’’ on Mt. Malinao accentuates the importance of this mountain as one of the hidden and neglected centers of endemism and biodiversity on the Bicol Peninsula. Previous conservation efforts on the peninsula have centered on the large and conspicuous Mt. Isarog (Goodman and Gonzales, 1990; Heaney et al., 1999; R. Brown et al., 2002b) and to a lesser extent, the picturesque volcano, Mt. Mayon. Other recent conservation and biodiversity survey efforts in the same biogeographic

region (R. Brown et al., 2002a; Diesmos et al., 2002b) have focused on non-montane habitats on Polillo Island (Hampson, 2001), Catanduanes Island (Auffenberg, 1988; Heaney et al., 1991; Ross and Gonzales, 1992), the Caramoan Peninsula (Auffenberg and Auffenberg, 1988, 1989), the volcanoes of southern Luzon (Custodio, 1986; Diesmos, 1998), and the southern Sierra Madre (R. Brown et al., 1999, 2000a, 2000b). While we do not wish to detract from these efforts, it is important to accentuate the importance of the lesser-known subcenters of Bicol biodiversity and to stress the role of amphibians as useful indicators of local centers of montane endemism and flagships for conservation (Diesmos, 1998; R. Brown et al., 2002a; Evans et al., 2003). As in other portions of the generic range of Platymantis (W. Brown et al., 1997b, 1999a; Alcala et al., 1998), the overall paucity of survey work at high elevations in the Philippines renders statements about species diversity in the islands premature. Despite recent work at montane sites throughout the archipelago, the Philippines suffers from a lack of comprehensive faunal inventories aimed at estimating amphibian diversity. We suspect that realistic estimates of Philippine amphibian diversity are attainable if government wildlife managers, university biologists, museum-based biodiversity specialists, and non-government conservation organizations personnel can work together to overcome bureaucratic obstacles to biodiversity studies in the Philippines. MATERIAL EXAMINED Platymantis dorsalis (13) Philippines, Negros Isl., Negros Oriental Prov., municipality of Valencia, 3.5 km S and 6.5 km W Valencia Town: FMNH 247742, 247746–48; Barangay Bongbong, Cuernos de Negros, Mt. Range, Mt. Talinis, Camp Lookout: FMNH 77890–02, 77727, TNHC 62095–99. Platymantis cagayanensis (8) Luzon Isl., Cagayan Province, Taggat Forest Reserve near Santa Praxedes Town: CAS 207447–451, PNM 6692–93, paratypes; PNM 6691, holotype. Platymantis corrugatus (14) Negros Isl., Negros Oriental Prov., Cuernos de Negros: FMNH 173174; ‘‘Canlaon Volcano:’’ FMNH 22508; municipality of Valencia, Barangay Bongbong, Camp Lookout, Cuernos de Negros Mt. Range, Mt. Talinis: TNHC 61972–76; Luzon Isl., Sorsogon Prov., municipality of Bulusan Barangay San Roque, Bulusan Volcano: FMNH 251646–47, 251649, 251651; Camarines Sur Prov., municipality of Naga City, Mt. Isarog: FMNH 25165–55, 251657. Platymantis indeprensus (14) Luzon Isl., Quezon

BROWN AND GONZALEZ—NEW PHILIPPINE PLATYMANTIS Isl., municipality of Tayabas, Baragany Lalo, Mt. Banahao, 1800 m: CAS 201013–14, 201178–83, 201185, 207452–54, PNM 6685–86, paratypes. Platymantis levigatus (7) Tablas Isl., Romblon Prov., municipality of San Agustin, Barangay Dubduban: CAS 136097, holotype; 137258–63, paratypes. Platymantis mimulus (10) Luzon Isl., Laguna Prov., municipality of Los Ban ˜ os, Barangay Batong Malake, Mt. Makiling: CAS 196362, holotype; TNHC 54930–34; University of the Philippines at Los Ban ˜ os campus, Mt. Makiling: FMNH 260355–56; Mt. Makiling: FMNH 110365; Los Ban ˜ os: FMNH 110372. Platymantis naomiae (4) Luzon Isl., Quezon Isl., municipality of Tayabas, Baragany Lalo, Mt. Banahao: CAS 204746, holotype; TNHC 62169–71. Platymantis pseudodorsalis (6) Luzon Isl., Quezon Isl., municipality of Tayabas, Baragany Lalo, Mt. Banahao: PNM 6689, holotype; 201536, 207456–59, paratypes. Platymantis pygmaeus (7) Luzon Isl., Isabela Prov., municipality of Palanan, Barangay Didian, Sitio Natapdukan: PNM 6255, holotype; CAS 204762–66, paratypes; Cagayan Prov., municipality of Claveria, Barangay Mabnang, Mabnang Falls: PNM 7523. Platymantis spelaeus (18) Negros Isl., Negros Oriental Prov., municipality of Basay, Tiyabanan Barrio: CAS 153470–83, paratypes; USNM 221838–39, FMNH 213331, paratypes. Platymantis taylori (20) Luzon Isl., Isabela Prov., municipality of Palanan, Barangay Didian, Sitio Natapdukan: PNM 6512–13, 6524, 6525–29, 6671, 6674–75, 6687–88, CAS 207440–207446, paratypes. Platymantis hazelae (17) Negros Isl., Negros Occidental Prov., ‘‘Mt. Canlaon:’’ FMNH 22512; municipality of Pula Pinto, SE slope Makawili Peak, Mt. Canloan: FMNH 95871–72; municipality of Valencia, Mt. Talinis: FMNH 247736, 247743–45, 250818; municipality of Valencia, Sitio Nasuji, Mt. Talinis Range, PNOCEDC Watershed: TNHC 62159–68; Platymantis isarog (8) Luzon Isl., Camarine Sur Prov., municipality of Naga City, Barangay Panicuason, Mt. Isarog National Park, Mt. Isarog CAS 197218, holotype; TNHC 61961–67. Platymantis lawtoni (5) Sibuyan Isl., Romblon Prov., E of Magdiwang, Mt. Guiting-guiting: FMNH 236092, 232096, 236142, 236146, 249701; Tablas Isl., Romblon Prov., municipality of San Agustin, Barangay Dubduban: CAS 135732, holotype. Platymantis montanus (13) Luzon Isl., Quezon Prov., municipality of Tayabas, Barangay Lalo, Mt. Banahao: TNHC 62149–58; CAS 200998–1000. Platymantis panayensis (17) Panay Isl., Aklan Prov., municipality of Libacao, Mt. Madja-as: PNM 2495, holotype; municipality of Nabas: CAS 137641– 42; Antique Prov., Mt. Madja-as, municipality of Culasi, Barangay Allojipan, ‘‘Hungud Tubig’’ Area: CMNH 4113–18, 4120, PNM 2314–20,

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paratypes. Platymantis polillensis (1) Polillo Isl., ‘‘Southern end of Polillo Island:’’ CAS 62250, holotype. Platymantis sierramadrensis (5) Luzon, Aurora Prov., municipality of San Luis; Dipiningan branch of the Cobatangan River drainage; 1.2 km S, 1.3 km E of Barangay Villa Aurora: PNM 6465, holotype, 5780, 5808; CMNH 5678– 79, 5904. Platymantis subterrestris (4) Luzon Isl., Mountain Prov., Mt. Data: CAS 204319–204322; ‘‘Lepanto Subprovince’’ CAS 61518, holotype. Platymantis guentheri (13) Mindanao Isl., Davao City Prov., ‘‘Tagum, Sitio Taglawig, near sea level:’’ FMNH 50571–73; Cotobato Prov., ‘‘Cotobato, near Milbuk:’’ FMNH 106656–57; Bohol Isl., Bohol Prov., municipality of Bilar, Barangay Logarito: RMB 2889, 2894–98, 2908, deposited at PNM; municipality of Carmen, Barangay Riverside: TNHC 56407. Platymantis banahao (14) Luzon Isl., Quezon Isl., municipality of Tayabas, Baragany Lalo, Mt. Banahao: CAS 201003–07, 201209–10, 201231, 201531–32, paratypes; TNHC 61968–71. Platymantis cornutus (2) Luzon Isl., Kalinga Prov., municipality of Balbalan, Barangay Balbalan: CAS 61476, holotype; CAS 231498. Platymantis insulatus (4) South Gigante Isl., Iloilo Prov., municipality of Estancia: CAS 117441, holotype; 119967–69, paratypes. Platymantis luzonensis (16) Luzon Isl., Laguna Prov., municipality of Los Ban ˜ os, Barangay Batong Malake, Mt. Makiling: CAS 196364, 196369–70, 200404–08, 210544–45, paratypes; Camarines Sur Prov., municipality of Naga City, Mt. Isarog: FMNH 251643–44, TNHC 62006–09. Platymantis negrosensis (12) Negros Isl., Negros Oriental Prov., Cuernos de Negros Mountain Range, Mt. Talinis, Lake Balinsasayo: CAS 137417–28, paratypes. Platymantis rabori (9) Leyte Isl., Leyte Prov., municipality of Abuera, Barangay Danao, Lake Danao: PNM 7776, 7693; Bohol Isl., Bohol Prov., municipality of Cantaub, Barangay Sierra Bullones: CAS 136888–96, paratypes. ACKNOWLEDGMENTS We thank the Protected Areas and Wildlife Bureau (PAWB) of the Philippine Department of Environment and Natural Resources (DENR) for facilitating collecting and export permits necessary for field portions of this and related studies. In particular, the guidance of C. Custodio was critical for obtaining access to sites, communicating with local authorities and landowners, and arranging research permits. We also thank the Mt. Mayon Protected Area Management Board, Tiwi and Malinao Municipal authorities (E. Kare, G. Gonzaga, S. Camu, R. Cruz, A. Can ˜ esa, N. Bringino, and S. Diaz), the Albay Provincial Council and Environmental Natural Resources Office (PENRO) office, Legaspi City PENRO,

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and National Power Corporation (especially C. Bandong) for logistical assistance. JCG thanks Alinao Guardian Mountaineers, A. Buizon, J. Corteza, J. Corteza, J. Corteza, A. Casin, J. Regalado, H. Austero, and L. Duran for field assistance. The assistance of V. Yngente, D. Yngente, M. Pedregosa, M. Ledesma, R. Cordial, E. Rico, B. Hernaez, A. and L. Dans, W. Reyes, R. Dumayas, and M. Valentin was also crucial for working on Mt. Malinao. Financial support for JCG’s fieldwork was provided by the American Zoo and Aquarium Association, the AZA Conservation Endowment Fund (especially W. Card), the Dallas Zoological Society (especially C. Bennett), the Cincinnati Zoo and Botanical Garden, Fauna and Flora International (especially W. Oliver), and the University of the Philippines at Los Ban ˜ os. Financial support for RMB’s fieldwork was provided by the Society of Systematic Biologists, the U.S. National Science Foundation, the Society for the Study of Amphibians and Reptiles, the American Society of Ichthyologists and Herpetologists, the Texas Memorial Museum, and the Section of Integrative Biology, University of Texas at Austin. For the loans of specimens and assistance while visiting museums, we thank J. Vindum and A. Leviton (CAS), R. Sison and A. Diesmos (PNM), J. Ferner (CMHN), A. Resetar and H. Voris (FMNH), R. Crombie (USNM), D. Cannatella and T. LaDuc (TNHC), L. Cardenas, E. Cosico, J. Lit, A. Cajano, and L. Afuang (UPLB-MNH). Financial support for RMB’s visits to CAS was provided by the Stearns Fellowship fund. We thank M. Garfield for illustrations and specimen photography, and D. and J. Esselstyn for help with the Tagalog abstract. Critical reviews of the manuscript were provided by J. Weghorst, C. Siler, and W. Duellman. We thank R. Fernandez and his family for their enthusiastic support, companionship in the field, and untiring efforts on our behalf during frequent visits to Albay Province. RMB thanks A. Diesmos for being his near-constant partner in the field for the past ten years, for help in all aspects of research in the Philippines, and for the steadfast example he has provided during countless nights of patiently waiting for frogs to call. LITERATURE CITED ALCALA, A. C., AND W. C. BROWN. 1998. Philippine Amphibians: an Illustrated Field Guide. Bookmark Inc., Makati City, Philippines. ALCALA, A. C., AND W. C. BROWN. 1999. Philippine frogs of the genus Platymantis (Amphibia: Ranidae). Philippine Journal of Science 128:281–287. ALCALA, A. C., W. C. BROWN, AND A. C. DIESMOS. 1998. Two new species of the genus Platymantis (Am-

phibia: Ranidae) from Luzon Island, Philippines. Proceedings of the California Academy of Sciences 50:381–388. ALLISON, A. 1996. Zoogeography of amphibians and reptiles of New Guinea and the Pacific region, p. 407–436. In: The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. A. Keast and S. E. Miller (eds.). SPB Academic Publishing, Amsterdam, The Netherlands. ALLISON, A., AND F. KRAUS. 2001. New species of Platymantis (Anura: Ranidae) from New Ireland. Copeia 2001:194–202. AUFFENBERG, W. 1988. Gray’s Monitor Lizard. University of Florida Press, Gainesville, Florida. AUFFENBERG, W., AND T. AUFFENBERG. 1988. Resource partitioning in a community of Philippine skinks (Sauria: Scincidae). Bulletin of the Florida State Museum of Biological Sciences 32:151–219. AUFFENBERG, W., AND T. AUFFENBERG. 1989. Reproductive patterns in sympatric Philippine skinks (Sauria: Scincidae). Bulletin of the Florida State Museum of Biological Sciences 34:201–247. BROWN, R. M., AND A. C. ALCALA. 2000. Geckos, cave frogs, and small land-bridge islands in the Visayan sea. Haring Ibon 2:19–22. BROWN, R. M., AND A. DIESMOS. 2002. (‘‘2001’’). Application of lineage-based species concepts to oceanic island frog populations: the effects of differing taxonomic philosophies on the estimation of Philippine biodiversity. Silliman Journal 42: 133–162. BROWN, R. M., A. DIESMOS, AND A. C. ALCALA. 2002a. (‘‘2001’’) The state of Philippine herpetology and the challenges for the next decade. Silliman Journal 42:18–87. BROWN, R. M., C. N. DOLINO, E. ALCALA, A. C. DIESMOS, AND A. C. ALCALA. 2003. (‘‘2002’’). The advertisement calls of two endangered species of endemic Philippine frogs: Platymantis spelaeus and P. insulatus (Anura; Ranidae). Silliman Journal 43: 91–109. BROWN, R. M., R. FERNANDEZ, C. RIVERO, R. BUENVIAJE, AND A. DIESMOS. 2002b. Mt. Isarog’s herpetological wonders. Haring Ibon 3:12–16. BROWN, R. M., J. A. MCGUIRE, AND A. C. DIESMOS. 2000a. Status of some Philippine frogs referred to Rana everetti (Anura: Ranidae), description of a new species, and resurrection of R. igorota Taylor 1922. Herpetologica 56:81–104. BROWN, R. M., J. A. MCGUIRE, J. W. FERNER, AND A. C. DIESMOS. 1999. A new species of diminutive scincid lizard (Squamata; Lygosominae; Sphenomorphus) from Luzon Island, Republic of the Philippines. Copeia 1999:362–370. BROWN, R. M., J. A. MCGUIRE, J. W. FERNER, N. ICARANGAL, AND R. S. KENNEDY. 2000b. Amphibians and reptiles of Luzon Island, II: preliminary report on the herpetofauna of Aurora Memorial National Park, Philippines. Hamadryad 25:175–195. BROWN, W. C. 1952. The amphibians of the Solomon islands. Bulletin of the Museum of Comparative Zoology 107:1–64.

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(RMB) NATURAL HISTORY MUSEUM, BIODIVERSITY RESEARCH CENTER, AND DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY, UNIVERSITY OF KANSAS, DYCHE HALL, 1345 JAYHAWK BOULEVARD, LAWRENCE, KANSAS 66045-7561; AND ( JCG) MUSEUM OF NATURAL HISTORY, UNIVERSITY OF ˜ OS, COLLEGE, LATHE PHILIPPINES AT LOS BAN GUNA, PHILIPPINES . E-mail: (RMB) [email protected]; and ( JCG) [email protected]. Send reprint requests to RMB. Submitted: 14 Aug. 2005. Accepted: 1 Dec. 2006. Section editor: T. W. Reeder.