Herpetologica, 65(1), 2009, 92–104 E 2009 by The Herpetologists’ League, Inc.
A NEW SPECIES OF LIMESTONE-FOREST FROG, GENUS PLATYMANTIS (AMPHIBIA: ANURA: CERATOBATRACHIDAE) FROM EASTERN SAMAR ISLAND, PHILIPPINES CAMERON D. SILER1,3,4, ANGEL C. ALCALA3, ARVIN C. DIESMOS2,3,
AND
RAFE M. BROWN1,3
1
Natural History Museum and Biodiversity Research Center, Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66045-7561, USA Herpetology Section, Zoology Division, Philippine National Museum, Rizal Park, Burgos St., Manila, Philippines 3 Angelo King Center for Research and Environmental Management, Silliman University, Dumaguete City, 6200, Philippines
2
ABSTRACT: A new species of forest frog of the genus Platymantis is described from an elevation of 140 m in the Taft Forest Reserve in eastern Samar Island, Philippines. It is assigned to the Platymantis guentheri Species Group, a group of primarily arboreal species, and is distinguished from these and other congeners by features of its external morphology and its preferred terrestrial, limestone microhabitat. Several striking morphological characters include a large body (34.2–39.1 mm SVL for 9 males and 44.3–49.8 mm SVL for 9 females), greatly expanded finger and toe discs, large eyes, spotted flanks, and sparsely-distributed, salmoncolored dorsal dermal tubercles. The new species represents the second largest Philippine Platymantis, the third terrestrial species in the P. guentheri Group, and the only known species of Platymantis from the Mindanao Faunal Region with a preference for forested, karst habitats. Key words: Biodiversity; Cryptic species; Endemism; Faunal Region; Limestone frogs; Platymantis guentheri Species Group; SW Pacific
morphology (W. C. Brown et al., 1997a,c; Alcala and W. C. Brown, 1999)—the P. dorsalis Group, the P. hazelae Group, and the P. guentheri Group. In addition to the morphological variation among Platymantis in the Philippines, the advertisement calls and preferred microhabitats of the species also vary (R. M. Brown et al., 2002; R. M. Brown and Gonzales, 2007). Surveys have shown that limestone-forest habitats of the Philippines possess highly distinctive species of Platymantis (W. C. Brown and Alcala, 1982; R. M. Brown and Alcala, 2000; Siler et al., 2007). In 1998 the late Walter C. Brown (see Alcala, 2004) informed RMB about a pair of highly distinctive Platymantis specimens from the Island of Samar that he had examined many years prior. He remarked that the suspected new species was ‘‘a giant form that looked like a cross between Platymantis speleaus and Platymantis insulatus.’’ Although Brown could not remember where the specimens were deposited, his verbal description of the suspected new species of giant Platymantis from Samar was quite specific and very detailed. Over the past decade we have been unable to locate the specimens in U.S. collections, but we have anxiously awaited the discovery of the new species that Brown
FROGS of the genus Platymantis have two geographic centers of diversity and endemism. One large group of species is situated in the Solomon Islands (W. C. Brown, 1952, 1997; Menzies, 2007) and nearby Bismarck (Zweifel, 1960, 1975; Foufopoulos and R. M. Brown, 2004; R. M. Brown et al., 2006a,b; Foufopoulos and Richards, 2007) and Admiralty archipelagos (26 species; Allison, 1996; Richards et al., 2007). Another large group of species occurs in the Philippines (27 species; Alcala and W. C. Brown, 1998, 1999; W. C. Brown et al., 1997a; R. M. Brown and Gonzales, 2007). Outside of these two large archipelagos, one species occurs in Palau (Crombie and Pregill, 1999); seven or eight are known from New Guinea faunal region (Zweifel, 1969; Allison, 1996; Gu¨nther, 1999, 2005; Menzies, 2007); two to four species occur in eastern Indonesia (Edgar and Lilley, 1993; Menzies, 1982a,b; including undescribed taxa; M. I. Setiadi and D. T. Iskandar, personal communication); and two others occur in Fiji (Gorham, 1965; Morrison, 2003). Within the Philippines, three species groups are recognized on the basis of external 4
CORRESPONDENCE: e-mail,
[email protected] 92
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FIG. 1.—Known occurrence of Platymantis bayani in eastern Samar Island, Philippines (black dot). The inset shows the location of Samar Island (colored black) within the Philippines. The three provinces of Samar Island are indicated by dashed lines.
predicted, ‘‘would eventually turn up on Samar.’’ During the month of October 2007, we conducted herpetological field surveys in the lower-elevation forest of the Municipality of Taft in eastern Samar Island (Fig. 1). Twentyone individuals of an undescribed species of Platymantis that match Walter Brown’s description were collected at an elevation of 140 m. Despite its striking tree frog appearance, the new species is found in terrestrial microhabitats in karst forest, an ecomorphological combination only shared with P. insulatus (R. M. Brown and Alcala, 2000). Herein, we describe the new species and report on its natural history, ecology, and habitat. MATERIALS AND METHODS We recorded morphometric data from alcohol-preserved specimens that were fixed in 10% formalin (Appendix 1). Sex was determined by gonadal inspection, and measurements were taken with digital calipers to the nearest 0.1 mm. To minimize inter-
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observer bias and other sources of potential error (Hayek et al., 2001; Lee, 1982), all measurements were scored by CDS. Descriptions of coloration were based on digital photographs and field notes of CDS. Characters measured follow the definitions of Siler et al. (2007) and include: snout–vent length (SVL), head length (HL), eye–narial distance (END), snout length (SNL), interorbital distance (IOD), internarial distance (IND), eye diameter (ED), horizontal tympanic annulus diameter (TAD), eye–tympanum distance (ETD), head width (HW), upper arm length (UAL), forearm length (FAL), tibia length (TBL), femur length (FEL), tarsus length (TSL), pes length (PL), manus length (ML), Toe IV length (Toe4L), Finger I length (Fin1L), Finger III length (Fin3L), Finger I disc width (Fin1DW), Finger III disc width (Fin3DW), Toe IV disc width (Toe4DW), and widths of penultimate phalanges of third finger (PpFin3) and the fourth toe (PpToe4). In the description, ranges are followed by mean 6 standard deviation in parentheses. Specimens were deposited at the Philippine National Museum (PNM) and Kansas University (KU). For this description, we follow Kraus and Allison (2007) in maintaining the masculine formations of species names within the genus Platymantis. RESULTS Platymantis bayani sp. nov. (Figs. 2–4) Holotype.—PNM 9501 (field no. CDS 2989, formerly KU 309260; Fig. 2), an adult male collected at 20:00 hr on 22 October 2007 at 140 m elevation in Taft Forest (11u 489 9.180 N, 125u 179 33.9360 E; WGS-84) Barangay San Rafael, Municipality of Taft, Eastern Samar Province, Samar Island, Philippines, by Cameron D. Siler and Jason B. Fernandez. Paratopotypes.—KU 309252–54, 309256, 309258–59, 309261, 309263–64, 309266–67, and 309269, PNM 9515–22, eleven males and nine females, collected between 14 and 23 October 2007. Diagnosis.—Platymantis bayani can be distinguished from other Philippine congeners
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FIG. 2.—Head of male holotype of Platymantis bayani (PNM 9501) in oblique view. Scale bar 5 5 mm.
by the following combination of characters: body size 34.2–39.1 mm SVL for males, 44.3– 49.8 mm SVL for females; digital disc of Finger III 2.7–4.03 width of penultimate phalanx in males and 2.8–4.53 in females; digital disc of Toe IV 2.1–3.33 width of penultimate phalanx in males and 2.9–3.73
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width in females (Fig. 3A,B); finger discs only slighter larger than toe discs; eye diameter large, 4.8–6.1 mm in adult males, and 5.4– 6.8 mm in adult females; skin granular on dorsal surfaces; dorsal dermal tubercles sparsely distributed, salmon-colored; supratympanic fold moderately protuberant; tympanum completely visible; dorsolateral stripes absent; and a preference for terrestrial microhabitat in karst forest. Mensural and diagnostic differences are provided in Tables 1 and 2. Comparisons.—The new species is assignable to the P. guentheri Group (P. banahao, P. cornutus, P. diesmosi, P. guentheri, P. insulatus, P. luzonensis, P. negrosensis, and P. rabori) based on the following combination of characters: digital discs greatly expanded and larger in fingers than in toes, first finger only slightly shorter than second, proximal portions of digits with narrow dermal folds in cross section, and subarticular tubercles large and highly protuberant (Fig. 3).
FIG. 3.—Ventral views of hand (A) and foot (B) of male holotype of Platymantis bayani (PNM 9501), hand (C) and foot (D) of male Platymantis insulatus (KU 300346), hand (E) and foot (F) of male Platymantis spelaeus (KU 300438), and hand (G) and foot (H) of male Platymantis paengi (KU 300209). Scale bar 5 2 mm.
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FIG. 4.—Photographs in life of Platymantis bayani paratopotypes exhibiting typical limestone-forest frog mottled color pattern and salmon-colored dorsal tubercles; (A) KU 309254, female, SVL 5 49.3 mm, and (B) KU 309252, male, SVL 5 34.7 mm. Photographs by C. Siler.
Samar Island 34.2–39.1 (37.1 6 1.8) 44.3–49.8 (46.4 6 2.1) 4.8–6.1 (5.5 6 0.5) 5.4–6.8 (6.1 6 0.4) 2.7–4.0 (3.3 6 0.4) 2.8–4.5 (3.9 6 0.5) 2.1–3.3 (2.8 6 0.4) 2.9–3.7 (3.1 6 0.3) 1.1–1.5 (1.3 6 0.1) 1.2–1.5 (1.3 6 0.1)
Supratympanic fold
Smooth, less prominent
2 No
+
+ + Cream
2
2 N/A
+ Dominant dorsal color 2 + + No Posterodorsal Posteroventral edge edge Smooth, not Rugose, prominent Rugose, prominent prominent
+
+
+
Luzon PAIC 24.1–32.5 (29.7 6 2.4) 32.6–33.8 (33.2 6 0.8) 2.7–3.9 (3.4 6 0.4) 3.6–3.9 (3.7 6 0.2) 2.2–3.3 (2.8 6 0.4) 3.0–3.2 (3.1 6 0.2) 1.6–2.2 (1.9 6 0.2) 1.8–1.9 (1.8 6 0.1) 1.6–2.0 (1.7 6 0.1) 2.0–2.1 (2.1 6 0.1)
luzonensis (9m, 2f)
Negros and Panay Islands 28.3–35.7 (31.1 6 4.0) 41.7–42.2 (42.0 6 0.3) 3.1–4.1 (3.7 6 0.6) 4.4–4.7 (4.6 6 0.2) 2.6–2.8 (2.7 6 0.1) 3.5–4.7 (4.1 6 0.8) 1.8–1.9 (1.8 6 0.1) 1.8–1.8 (1.8 6 0.0) 1.8–2.0 (1.9 6 0.1) 1.7–2.0 (1.9 6 0.3)
negrosensis (3m, 2f)
Gigantes Islands 34.3–38.0 (36.7 6 1.3) 40.2–42.9 (41.8 6 1.0) 3.7–4.9 (4.2 6 0.4) 5.1–5.7 (5.3 6 0.2) 2.7–3.1 (2.9 6 0.1) 2.3–2.8 (2.7 6 0.2) 1.9–3.4 (2.6 6 0.4) 2.0–2.6 (2.3 6 0.3) 1.1–1.5 (1.3 6 0.1) 1.2–1.6 (1.4 6 0.1)
insulatus (8m, 6f)
+
+, moderate
Mindanao PAIC 26.5–29.8 (28.6 6 1.1) 32.3–37.3 (34.6 6 2.0) 2.8–3.6 (3.3 6 0.3) 3.7–4.5 (3.9 6 0.3) 2.4–5.2 (3.2 6 0.8) 2.6–5.7 (4.0 6 1.2) 1.5–3.5 (2.4 6 0.6) 2.3–4.3 (3.2 6 0.7) 1.2–1.7 (1.4 6 0.1) 1.1–1.7 (1.4 6 0.2)
guentheri (11m, 8f)
+, dense + Dominant Dominant dorsal color dorsal color + + Posterior Posteroventral edge edge Rugose, Smooth, less prominent prominent
+
+
Mindanao PAIC 30.4–33.5 (32.0 6 2.2) 36.4–44.4 (41.5 6 3.6) 3.8–3.9 (3.9 6 0.1) 4.0–5.7 (4.9 6 0.7) 2.8–3.5 (3.2 6 0.5) 3.3–4.2 (3.6 6 0.3) 2.0–2.7 (2.3 6 0.5) 1.6–2.5 (2.3 6 0.3) 1.3–1.6 (1.4 6 0.2) 1.4–1.9 (1.6 6 0.2)
rabori (2m, 6f)
1.9–2.1 (2.0 6 0.1) N/A
1.8–1.9 (1.8 6 0.0) N/A
+ Dominant dorsal color 2 Posterodorsal edge Smooth, less prominent
2 to granular
Smooth, not prominent
2 No
2 N/A
2
+
1.6–2.0 (1.7 6 0.2) N/A
1.9–2.3 (2.0 6 0.2) N/A
+
2.5–2.9 (2.7 6 0.2) N/A
3.0–4.2 (3.5 6 0.6) N/A
N. Luzon Island 26.3–30.4 (27.8 6 2.2) N/A
cornutus (3m)
2.8–3.2 (3.0 6 0.2) N/A
3.6–4.1 (3.8 6 0.2) N/A
SE Luzon Island 32.4–36.3 (33.9 6 1.7) N/A
banahao (4m)
2 Posterodorsal edge Smooth, less prominent
2 N/A
2 to granular
2
SE Luzon Island 28.5–39.2 (35.3 6 3.6) 49.0–52.6 (50.8 6 2.6) 3.7–5.4 (4.7 6 0.5) 5.9–6.1 (6.0 6 0.1) 1.5–2.1 (1.7 6 0.2) 1.6–1.7 (1.6 6 0.1) 1.2–2.2 (1.7 6 0.4) 1.7–2.1 (1.9 6 0.3) 0.9–1.4 (1.1 6 0.2) 0.8–0.8 (0.8 6 0.0)
diesmosi (8m, 2f)
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Dorsal folds/ridges Tympanum hidden
Fin3DW/PpFin3 (m) Fin3DW/PpFin3 (f) Toe4DW/PpToe4 (m) Toe4DW/PpToe4 (f) Fin3DW/Toe4DW (m) Fin3DW/Toe4DW (f) Greatly expanded digital disks + Dorsal skin rugosity 2 to granular Dorsal dermal tubercles +, sparse Tubercle color Salmon
ED (f)
ED (m)
SVL (f)
SVL (m)
Range
bayani sp. nov. (9m, 9f)
TABLE 1.—Summary of qualitative diagnostic characters (present 5 +, absent 5 2) in Platymantis bayani and specimens of other members of the P. guentheri Species Group. Sample size for each sex, body size, eye diameter, digit disk proportions, and general geographical distribution (PAIC 5 Pleistocene Aggregate Island Complexes, sensu Brown and Diesmos, 2002) are included for reference (SVL and ED given as range over mean 6 standard deviation; digit disk proportions given as percentage over mean 6 standard deviation). Only adult specimens in good preservation condition were used for morphometric analyses.
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+ 2 +, poorly defined Arboreal 2 2 + or 2, faint Arboreal + or 2 2 +, well defined Shrub/Arboreal + or 2 2 +, well defined Arboreal + or 2 2 +, well defined Arboreal + 2 +, well defined Arboreal Ground; karst habitat Microhabitat preference
2 + + or 2, poorly defined Ground; karst habitat 2 + +, well defined Dorsolateral stripes Spotted flanks Limb bands
cornutus (3m) banahao (4m) guentheri (11m, 8f) rabori (2m, 6f) luzonensis (9m, 2f) negrosensis (3m, 2f) insulatus (8m, 6f) bayani sp. nov. (9m, 9f)
TABLE 1.—Continued.
2 2 +, well defined Ground; cliff edges
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diesmosi (8m, 2f)
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From all members of the P. dorsalis Group (P. cagayensis, P. corrugatus, P. dorsalis, P. indeprensus, P. levigatus, P. mimulus, P. naomiae, P. paengi, P. pseudodorsalis, P. pygmaeus, P. spelaeus, and P. taylori), Platymantis bayani is distinguished by the presence (vs. absence) of the following features: greatly expanded digital discs (Fig. 3); slightly larger finger discs than toe discs; first finger shorter than second finger (Fig. 3); and sparsely distributed, salmon-colored, dorsal dermal tubercles (vs. presence and dominant dorsal color in P. dorsalis, P. paengi, P. spelaeus, and P. taylori, and absence in P. cagayensis, P. corrugatus, P. indeprensus, P. levigatus, P. mimulus, P. naomiae, P. pseudodorsalis, and P. pygmaeus; Table 2). Platymantis bayani is further distinguished from all members of the P. dorsalis Group, except P. spelaeus, by having a much larger body size, and from all members except P. paengi and P. spelaeus, by its microhabitat preference (terrestrial substrates in limestone forest vs. leaf litter of forest floor and/or herb-layer vegetation in P. cagayensis, P. corrugatus, P. dorsalis, P. indeprensus, P. mimulus, P. naomiae, P. pseudodorsalis, P. pygmaeus and P. taylori, or river banks and rocks in P. levigatus). From P. cagayensis, P. corrugatus, P. dorsalis, P. indeprensus, P. mimulus, P. naomiae, P. pygmaeus and P. taylori, the new species is distinguished by the absence (vs. presence) of dorsal folds and ridges. The new species is distinguished from all P. hazelae Group species (P. hazelae, P. isarog, P. lawtoni, P. montanus, P. panayensis, P. polillensis, P. sierramadrensis, P. subterrestris) by its larger body size; the first finger much shorter than the second; narrow dermal phalanges on the digits; finger discs only slightly larger than toe disks; protuberant (vs. flattened) subarticular tubercles, and a preferred microhabitat of terrestrial substrates in limestone forest (vs. arboreal habitat, and/or shrub-layer vegetation). In the following section, the new species is compared with all species of the P. guentheri Group. The character states for congeners can be found in Tables 1 and 2. The new species differs from all other species of the P. guentheri Group, except for P. diesmosi, by its larger body size and greater
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TABLE 2.—Summary of mensural characters (in mm) and selected qualitative diagnostic characters (present, absent) in Platymantis bayani and other Philippine limestone forest frog species. Sample size, body size, species group, and general geographical distribution (PAIC 5 Pleistocene Aggregate Island Complexes, sensu Brown and Diesmos, 2002) are included for reference (SVL given as range over mean 6 standard deviation).
SVL (m) SVL (f) Range Species Group Digital disks Dorsal skin rugosity Dorsal dermal tubercles Flanks
bayani sp. nov. (9m, 9f)
insulatus (8m, 6f)
spelaeus (4m, 3f)
paengi (15m)
34.2–39.1 44.3–49.8 Samar Island guentheri Widely expanded 2 to granular +, sparse Spotted
34.3–38.0 40.2–42.9 Gigantes Islands guentheri Widely expanded 2 2 Spotted
37.1–44.3 53.5–57.5 SW Negros Island dorsalis Non-expanded +, 2 +, small Immaculate
27.7–34.3 N/A NW Panay Island dorsalis Non-expanded + + Spotted
eye diameter. The new species differs from P. negrosensis, P. luzonensis, P. rabori, and P. guentheri by lacking rugose dorsal skin, dorsal folds and ridges, and dorsolateral stripes. It is distinguished from P. diesmosi, P. negrosensis, P. luzonensis, P. rabori, P. guentheri, and P. banahao by having a completely visible tympanum, dark brown to black spots on the flanks, sparsely distributed dorsal dermal tubercles that are salmon-colored, and a preference for terrestrial microhabitats in limestone forest. Platymantis bayani is further distinguished from P. negrosensis, P. luzonensis, and P. rabori by having a smooth and less prominent supratympanic fold. From P. cornutus, P. negrosensis, P. luzonensis, and P. banahao, it differs in having slightly larger finger discs than toe discs. The new species differs from P. diesmosi in having greatly expanded digital discs, and from P. cornutus, by having dark brown to black spots on the flanks, sparsely distributed, salmon-colored, dorsal dermal tubercles, a slightly prominent supratympanic fold, well-defined limb bands, the absence of dorsolateral stripes, and a preference for terrestrial microhabitats in limestone forest. From P. insulatus, the new species is distinguished by the presence of dorsal dermal tubercles and a slightly prominent supratympanic fold. Finally, P. bayani differs from P. insulatus and P. banahao in having a distinct, transverse limb-banding pattern. Description of holotype.—A mature male; habitus robust; head broader than body, HL 37.7% SVL, 91.2% HW; snout protruding moderately beyond lower jaw, snout tip comes to sharp point in dorsal and lateral aspects;
eyes large and protruding laterally, ED 15.6% SVL, 41.3% HL; lips slightly flared and swollen (Fig. 2); interorbital region flat; ED 137.0% SNL, 1.73 END; pupil horizontally elliptical; canthus rostralis laterally concave in dorsal aspect; loreal region greatly concave, sloping ventrolaterally to labium; nostrils oriented posterolaterally, protruding laterally to sharp points from snout tip; END 2.33 SNL; internarial region flat; tympanic annulus distinct, its diameter 0.53 ED; supratympanic fold moderately protuberant, extending from posterior corner of eye across dorsal margin of tympanum to supra-axillary region. Tongue elongate, wider posteriorly, with deep posterior notch and narrow anterior attachment; choanae situated at anterolateral edge of palate, oval, widely separated by distance three to four times greater than diameter of single choana; dentigerous process of vomer distinct, with two teeth located on anterior 1/3 of process; dentigerous process angled slightly anterolaterally, approximately at 10–20u incline with closest (posterior) points separated by distance equal to 2.03 diameter of single choana, their most distant (anterior) ends separated by distance equal to 3.03 diameter of single choana; short vocal slits at posteroventral margin of mouth. Skin of dorsal surfaces of trunk and head coarsely granular, distinctly shagreen; irregular, moderately protuberant tubercles sparsely distributed across dorsal surfaces of body and limbs; ventral surfaces of head smooth; lateral and ventral surfaces of limbs finely granular with dorsal surfaces coarsely granular; flanks shagreen. Manus length 58.0% pes length; tibia length 108.8% pes length; tibia length 54.6% SVL;
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fingers slender, long; terminal discs greatly expanded (Fig. 3A) with pronounced distal, circummarginal groove; dorsal surfaces of terminal phalanges with small, cutaneous supra-articular flaps between ultimate and penultimate phalanges; relative lengths of fingers I 5 II , IV , III; subarticular tubercles prominent, convex, granular in texture; one subarticular tubercle below Digits I and II, two tubercles under Digits III and IV; supernumerary tubercles distinct, round, prominent; present at bases of Digits I–IV; inner metacarpal tubercle enlarged, elongate, crescent shaped; outer metacarpal tubercle partially divided into oval, slightly raised preaxial tubercle of moderate size, and round postaxial tubercle equal in size to preaxial metacarpal tubercle; nuptial pads absent; forearms slender, not hypertrophied in males. Tarsus smooth, lacking folds, flaps, or tubercles; terminal discs of toes greatly expanded, with circummarginal grooves; supra-articular cutaneous flaps above ultimatepenultimate phalangeal articulations; plantar surface of foot smooth, with well-developed, prominently rounded subarticular tubercles (Fig. 3B); supernumerary tubercles absent; relative lengths of toes I , II , V , III , IV; outer metatarsal tubercle small, round; inner metatarsal tubercle prominent, with sharp inner edge, length equal to distance from base of Toe I to first subarticular tubercle; toes with vestige of basal interphalangeal web, nearly extending to the level of the proximal subarticular tubercles between Toes III and IV. Coloration of holotype in preservative.— Dominant dorsal color pattern on head and body mottled with gray and silver blotches and spots interspersed throughout a dominant background of black and dark brown; dorsal surfaces of limbs banded with alternating green-brown and dark brown transverse bars; interorbital bar slightly darker than dominant dorsal color pattern; snout region slightly darker than dorsum, and nearly uniform black; wide black bar extending from snout tip, across canthus rostralis to anterior edge of eye; a dark blotch covering anteroventral edge of tympanum, ventral two thirds of tympanum lighter gray; temporal region dorsally mottled
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dark brown and black; lips lighter gray with faint vertical dark brown labial bars and speckles, bordered anteriorly by lighter brown lower edge of canthus and subocular regions; transverse, dark brown bars on limbs similar in color to dorsum, coalescing into mottled patterns on elbows and knees; four bars on forearms, four on thigh, four on tibia, three on tarsus; spotted flanks, ground color lighter grayish tan with dark brown blotches and spots, blending into the darker dorsal pattern (above) and the lighter ventral pattern (below); dorsal surfaces of manus, pes, and digits light grayish tan with dark brown blotches above each joint. Throat marbled light cream to tan with speckled green-brown blotches; periphery of throat and lower lips lighter with anterior edge of lower lip possessing outline of light cream spots; chest light cream with less dense speckled green-brown blotches; lighter ventral coloration limited to small area on ventral surface of forelimb near limb insertion; ventral surface of upper- and forearms darker cream with green-brown blotches; venter light cream with nearly indistinct light green-brown blotches that decrease in size posteriorly until blending with uniformly light cream groin; ventral surfaces of thigh uniformly mottled cream and green-brown; ventral surface of tibia and tarsus mottled green-brown, similar to forearms; palmar and plantar surfaces of hands and feet dark green-brown with light cream subarticular tubercles; iris pale gray above and dark gray below pupil. Coloration of holotype in life.—Dorsum and head mottled light green, dark green, and black (Fig. 4); interorbital bar distinct, black, and bordered anteriorly by lighter, green-brown border; dorsal surfaces of limbs with alternating dark brown and green transverse bars; dorsal surface of body with low, salmon-colored tubercles; series of dark brown to black blotches and spots across flanks; pupil bordered by bright gold ciliary ring of iris, blending into the predominant mottled golden-brown and dark brown color pattern of iris. Measurements of holotype (mm).—SVL 39.1; HL 14.7; HW 16.2; SNL 4.4; IND 3.6; IOD 3.2; ED 6.1; END 3.5; TAD 2.8; ETD 1.0; UAL 7.4; FAL 9.0; FEL 20.1; TBL 21.3;
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TABLE 3.—Summary of univariate morphological variation among mensural characters in the type series of Platymantis bayani. See text for definitions of abbreviations. Female n 5 9
SVL HL END SNL IOD IND ED TAD ETD HW UA FAL FL TBL TSL PL ML Toe4L Fin1L Fin3L Fin1DW Fin3DW Toe4DW PpFin3 PpToe4
44.3–49.8 16.2–18.8 3.3–4.8 3.8–6.1 4.0–4.9 3.6–4.3 5.4–6.8 2.3–3.3 1.1–1.7 18.4–20.6 8.2–9.9 12.0–13.1 23.7–26.2 21.0–26.9 14.7–15.9 22.9–25.3 13.5–14.8 21.8–23.9 7.2–8.4 12.7–13.9 1.6–2.0 2.3–2.9 1.7–2.3 0.6–0.8 0.6–0.7
(46.4 6 2.1) (17.3 6 0.8) (4.1 6 0.5) (5.3 6 0.8) (4.3 6 0.3) (3.9 6 0.2) (6.1 6 0.4) (2.8 6 0.3) (1.4 6 0.2) (19.7 6 0.7) (9.1 6 0.6) (12.6 6 0.4) (24.9 6 0.9) (25.8 6 1.8) (15.1 6 0.4) (23.9 6 0.9) (14.2 6 0.3) (22.8 6 0.6) (7.9 6 0.4) (13.2 6 0.4) (1.8 6 0.1) (2.7 6 0.2) (2.0 6 0.2) (0.7 6 0.1) (0.6 6 0.1)
TSL 12.8; ML 11.4; PL 19.6; Fin1L 6.1; Fin3L 10.3; Toe4L 18.2; Fin3DW 1.8; Toe4DW 1.5; PpFin3 0.7; PpToe4 0.7. Variation.—Summaries of univariate morphological variation in the series are presented in Table 3. Dominant color pattern consistently mottled in all observed specimens; however, the degree of contrast between the dominant ground color and dark blotches varies among individuals, with two patterns of contrast observed. The first pattern consists of dark brown to black blotches intermixed with a light cream to green ground color (KU 309253–54, 309256, 309258–59, 309261, and 309264). The second pattern was consists of dark green blotches intermixed with the same light cream to green ground color (KU 309266, 309269, 309252, 309263, and 309267). Distribution.—Platymantis bayani is known only from the type locality (Fig. 1). Etymology.—The name of the new species is derived from the Tagalog (Filipino) term ‘‘bayani,’’ meaning ‘‘highly respected,’’ ‘‘heroic,’’ or ‘‘hero.’’ We are pleased to name this
Male n 5 9
34.2–39.1 14.1–15.5 3.2–4.4 4.0–5.9 3.0–4.2 2.9–3.6 4.8–6.1 1.9–2.8 1.0–1.6 15.1–17.0 7.3–8.1 9.8–10.6 19.6–20.7 21.2–22.8 12.0–13.2 18.7–20.3 11.2–12.3 18.1–20.0 6.1–7.1 10.3–11.5 1.3–1.6 1.8–2.5 1.5–1.8 0.5–0.7 0.5–0.7
(37.1 6 1.8) (14.8 6 0.5) (3.9 6 0.4) (5.0 6 0.8) (3.5 6 0.3) (3.2 6 0.2) (5.5 6 0.5) (2.4 6 0.3) (1.3 6 0.2) (15.9 6 0.6) (7.6 6 0.3) (10.3 6 0.2) (20.2 6 0.4) (21.7 6 0.5) (12.6 6 0.4) (19.7 6 0.6) (11.7 6 0.4) (18.8 6 0.6) (6.4 6 0.4) (10.8 6 0.4) (1.4 6 0.1) (2.1 6 0.2) (1.6 6 0.1) (0.7 6 0.1) (0.6 6 0.1)
new species for the late Walter C. Brown in recognition of his numerous contributions to Philippine herpetology and in acknowledgment of his lifelong commitment to the study of biodiversity in the SW Pacific. Suggested common name: Walter’s Limestone Forest Frog. Ecology and natural history.—Platymantis bayani occurs in primary- and secondarygrowth forest on karst substrate. The advertisement call of the species is unknown. Both males and females were found perched on top of limestone rocks within large rock formations in the forest; they never were observed perching in trees or shrubs as do most species in the P. guentheri Group (R. M. Brown and Gonzales, 2007). Platymantis bayani was well camouflaged against the limestone (much like P. insulatus, P. speleaus, and P. paengi; Fig. 4), and collectors found it difficult to locate and capture individuals against the limestone substrate. When disturbed, the new species immediately jumped back into the rocks and escaped into limestone crevices. No egg
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masses or juveniles were found; other species of Platymantis have direct development (W. C. Brown and Alcala, 1982a,b). Platymantis spelaeus has been observed by one of us (CDS) to deposit direct-developing eggs in limestone crevices and the same is suspected for P. insulatus (R. M. Brown and Alcala, 2000). We assume that this species also breeds in limestone crevices and small caves (R. M. Brown and Alcala, 2000; R. M. Brown et al., 2003). Sympatric anuran species include Kalophrynus pleurostigma, Limnonectes magnus, Megophrys stejnegeri, Platymantis cf. dorsalis, P. cf. corrugatus, P. rabori, P. guentheri, Polypedates leucomystax, Philautus leitensis, Rhacophorus bimaculatus, R. appendiculatus, R. pardalis, Oreophryne sp., Rana grandocula, R. albotuberculata, and Occidozyga laevis. DISCUSSION With the new species described here, a total of 28 Philippine species of Platymantis are now recognized. The species total for the country has increased dramatically over the last several decades; seven species were recognized between 1950 and 1970 (Inger, 1954; W. C. Brown and Alcala, 1970a,b) and 12 by the late 1980’s (W. C. Brown and Alcala, 1982a). An additional dozen species were described in the late 1990’s (W. C. Brown et al., 1997a,b,c, 1999), several more species have been described recently (R. M. Brown and Gonzales, 2007; Siler et al., 2007), and numerous unnamed species await description (Brown, 2004; Brown et al., 2008). Within the Philippine radiation, high levels of cryptic diversity (Alcala et al., 1998; W. C. Brown et al., 1997b, 1999) has only recently been uncovered (Brown, 2004; Brown et al., 2008) due to attention on advertisement-call diversity, microhabitat preferences, and gene sequence data Platymantis bayani is the fourth Philippine limestone frog to be described, the third terrestrial member of the P. guentheri Group (the others being P. insulatus and P. diesmosi), and the only limestone species known to occur in the Mindanao Faunal Region. Platymantis spelaeus from SW Negros Island, P. insulatus from the Gigantes Islands, and P. paengi from NW Panay Island are the three previously
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known karst-forest frogs found in the Visayan (Central) Philippine Islands. All four species seem to be obligate inhabitants of limestone forests. Although potentially restricted in geographic ranges, the species are quite common in the rainy season, but can seem to be rare in times of high temperature and aridity when we assume they retreat into subterranean limestone caves and crevices (R. M. Brown and Alcala, 2000; R. M. Brown et al., 2003). The recent discoveries of new species of Philippine frogs and gekkos with preferences for limestone forests (Siler et al., 2007; Roesler et al., 2006) highlight the degree to which this imperiled habitat type (Clements et al., 2006) is poorly known. Future survey efforts should make a point to target limestone forest communities. Our general impression is that these habitat types have been overlooked in the past because surrounding forests are dry, stunted, difficult to sample, and viewed by field biologists as non-productive. The heavily mottled, predominant color pattern of Platymantis bayani resembles P. spelaeus, P. insulatus, and P. paengi. The conspicuous similarity of color patterns among these taxa suggests to us the possibility of convergent selection on color pattern in limestone habitats. It is unclear as to which species P. bayani is most closely related, but phenotypic similarity with P. insulatus is striking. The new species was quite common at the type locality, and it may posses a wider geographical distribution, possibly throughout the northern islands of the Mindanao Faunal Region (Samar, Leyte, Biliran, Mindanao, Dinagat, Siargao, and smaller associated islands). It is possible that a primary limiting factor to the range of P. bayani is the presence of relatively undisturbed limestone formations and forest cover. We note that both P. speleaus and P. insulatus persist quite well, despite the near complete removal of forest on the Gigante Island group (P. insulatus) and southern Negros Island (P. spelaeus). Intensive survey efforts will be required to fully document the distribution of the new species and arrive at a reasonable assessment of its conservation status. At present, we consider the status of the new species to be ‘‘data
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deficient’’ pending the collection of much needed data on its distribution and habitat requirements. 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 this and related studies, wherein we are particularly grateful to M. Lim, C. Custodio, and A. Tagtag. We also thank the Arnie Viojan and the Regional Office of the Protected Areas, Wildlife, and Coastal Zones Management Services (PAWCZMS), and Director Manolito D. Ragub, Eirish Mate, and the headquarters of the Samar Island Natural Parks (SINP), whose logistical support and guidance were instrumental throughout our visit to Samar. Jason Fernandez assisted in the field. Financial support for fieldwork was provided by a Panorama Fund grant to CDS from The University of Kansas Natural History Museum and Biodiversity Institute, a Madison and Lila Self Fellowship to CDS from the University of Kansas, NSF EF-0334952 to L. Trueb and R. Brown and DEB 0743491 to RMB. For the loans of specimens (museum abbreviations follow Leviton et al., 1985), we thank J. Vindum and A. Leviton (CAS), R. Sison and A. C. Diesmos (PNM), J. Ferner (CMNH), A. Resetar and H. Voris (FMNH), R. Crombie (USNM), and T. LaDuc (TNHC). Critical reviews of the manuscript were provided by L. Trueb and J. Esselstyn. We thank Michael Garfield for illustrating the holotype for Figure 2. CDS thanks the Philippine-American Education Foundation, A.C. Alcala and family, and M. Diesmos for their continued support, as well as CAS’s Stearns Fellowship for funding a recent visit to examine comparative material. Finally, we owe a debt of gratitude to the late Walter C. Brown who alerted RMB to the presence of this new species on Samar more than 10 yr before we were able to find it. Once again, Walter was right.
LITERATURE CITED ALCALA, A. C. 2004. A life of service to science: a tribute to Walter Creighton Brown, 1913–2002. Silliman Journal 45:258–262. 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 (Amphibia: 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. Pp. 407–436. In A. Keast and S. E. Miller (Eds.). The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Publishing, Amsterdam, The Netherlands. BROWN, R. M. 2004. Evolution of Ecomorphological Variation and Acoustic Diversity in Mate-Recognition
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Signals of Southeast Asian Forest Frogs (Subfamily Platymantinae). Doctoral dissertation, University of Texas at Austin, Austin, Texas, U.S.A. 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 J. C. GONZALES. 2007. A new forest frog of the genus Platymantis (Amphibia; Anura; Ranidae) from the Bicol Peninsula of Luzon Island, Philippines. Copeia 2007:251–266. BROWN, R. M., A. DIESMOS, AND A. C. ALCALA. 2002. The state of Philippine herpetology and the challenges for the next decade. Haring Ibon 42:18–87. BROWN, R. M., A. C. DIESMOS, AND A. C. ALCALA. 2008. Philippine amphibian biodiversity is increasing in leaps and bounds. Pp. 82–83. In S. N. Stuart, M. Hoffmann, J. S. Chanson, N. A. Cox, R. Berridge, P. Ramani, and B. E. Young (Eds.). Threatened Amphibians of the World. Lynx Ediciones, Barcelona, Spain; IUCN—The World Conservation Union, Gland, Switzerland; and Conservation International, Arlington, Virginia, U.S.A. BROWN, R. M., C. N. DOLINO, E. ALCALA, A. C. DIESMOS, AND A. C. ALCALA. 2003. The advertisement calls of two endangered species of endemic Philippine frogs: Platymantis spelaeus and P. insulatus (Anura; Ranidae). Haring Ibon 43:91–109. BROWN, R. M., J. FOUFOPOULOS, AND S. J. RICHARDS. 2006. New species of Platymantis (Amphibia; Anura: Ranidae) from New Britain and redescription of the poorly known Platymantis nexipus. Copeia 2006:674–695. BROWN, R. M., S. J. RICHARDS, J. SUKUMARAN, AND J. FOUFOPOULOS. 2006. A new morphologically cryptic species of forest frog (genus Platymantis) from New Britain Island, Bismarck Archipelago. Zootaxa 1334:45–68. BROWN, W. C. 1952. The amphibians of the Solomon Islands. Bulletin of the Museum of Comparative Zoology, Harvard 107:1–64. BROWN, W. C. 1997. Biogeography of amphibians in the islands of the southwest pacific. Proceedings of the California Academy of Sciences 50:21–38. BROWN, W. C., AND A. C. ALCALA. 1970a. A new species of the genus Platymantis (Ranidae) with a list of the amphibians known from South Gigante Island, Philippines. Occasional Papers of the California Academy of Sciences 84:1–7. BROWN, W. C., AND A. C. ALCALA. 1970b. The zoogeography of the Philippine Islands, a fringing archipelago. Proceedings of the California Academy of Sciences 38:105–130. BROWN, W. C., AND A. C. ALCALA. 1982a. A new cave Platymantis (Amphibia: Ranidae) from the Philippine Islands. Proceedings of the Biological Society of Washington 95:386–391. BROWN, W. C., AND A. C. ALCALA. 1982b. Modes of reproduction of Philippine anurans. Pp. 416–428. In A. G. J. Rodin and K. Miyata (Eds.). Advances in Herpetology and Evolutionary Biology. Museum of Comparative Biology, Cambridge, Massachusetts, U.S.A. BROWN, W. C., R. M. BROWN, AND A. C. ALCALA. 1997a. Species of the hazelae group of Platymantis from the Philippines, with descriptions of two new species.
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Proceedings of the California Academy of Sciences 49:405–421. BROWN, W. C., A. C. ALCALA, AND A. C. DIESMOS. 1997b. A new species of the genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines. Proceedings of the Biological Society of Washington 110:18–23. BROWN, W. C., A. C. ALCALA, A. C. DIESMOS, AND E. ALCALA. 1997c. Species of the guentheri group of Platymantis from the Philippines with descriptions of four new species. Proceedings of the California Academy of Sciences 50:1–20. BROWN, W. C., A. C. ALCALA, AND A. C. DIESMOS. 1999. Four new species of the genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines. Proceedings of the California Academy of Sciences 51:449–460. CLEMENTS, R., N. SODHI, M. SCHILTHUIZEN, AND P. K. L. NG. 2006. Limestone karsts of Southeast Asia: imperiled arks of biodiversity. Bioscience 56:733–742. CROMBIE, R. I., AND G. K. PREGILL. 1999. A checklist of the herpetofauna of the Palau Islands (Republic of Belau), Oceania. Herpetological Monographs 13:29–80. EDGAR, P. W., AND R. P. H. LILLEY. 1993. Herpetofauna survey of Manusela National Park. Pp. 131–141. In I. D. Edwards, A. A. Macdonald, and J. Proctor (Eds.). Natural history of Seram, Maluku, Indonesia. Intercept Ltd, Andover, Massachusetts, U.S.A. FOUFOPOULOS, J., AND R. M. BROWN. 2004. A new frog of the genus Platymantis (Amphibia; Anura; Ranidae) from New Britain, with a redescription of the poorly known Platymantis macrosceles. Copeia 2004:825–841. FOUFOPOULOS, J., AND S. RICHARDS. 2007. Amphibians and reptiles of New Britain Island, Papua New Guinea: diversity and conservation status. Hamadryad 31:176–201. GORHAM, S. W. 1965. Fiji frogs, with synopses of the genera Cornufer and Platymantis. Dunker and Humbolt, Berlin, Germany. GU¨NTHER, R. 1999. Morphological and bioacoustic characteristics of frogs of the genus Platymantis (Amphibia: Ranidae) in Irian Jaya, with descriptions of two new species. Mitteilungen aus dem Museum fu¨r Naturkunde in Berlin-Zoologische Reihe 75:317–335. GU¨NTHER, R. 2005. A new species of the frog genus Platymantis from the mountains of Yapen Island, northern Papua Province, Indonesia (Amphibia: Anura: Ranidae). Zoologische Abhandlungen 55:85–94. HAYEK, L.-A. C., W. R. HEYER, AND C. GASCON. 2001. Frog morphometrics: a cautionary tale. Alytes 18:153–177. INGER, R. F. 1954. Systematics and zoogeography of Philippine Amphibia. Fieldiana: Zoologica 33:182–531. KRAUS, F., AND A. ALLISON. 2007. Two new species of Platymantis (Anura: Ranidae) from New Britain. Zootaxa 1485:13–32. LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802–821. LEE, J. C. 1982. Accuracy and precision in anuran morphometrics: artifacts of preservation. Systematic Zoology 31:266–281. MENZIES, J. I. 1982a. The voices of some male Platymantis species of the New Guinea region. British Journal of Herpetology 6:1–5.
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MENZIES, J. I. 1982b. Systematics of Platymantis papuensis (Amphibia: Ranidae) and related species in the New Guinea region. British Journal of Herpetology 6:236–245. MENZIES, J. I. 2007. The frogs of New Guinea and the Solomon Islands. Pensoft Publishing, Sophia, Bulgaria. MORRISON, C. 2003. A field guide to the herpetofauna of Fiji. Institute of Applied Sciences, University of the South Pacific, Suva, Fiji. RICHARDS, S. J., A. L. MACK, AND C. C. AUSTIN. 2007. Two new species of Platymantis (Anura: Ceratobatrachidae) from the Admiralty Archipelago, Papua New Guinea. Zootaxa 1639:41–55. ROSLER, H., C. D. SILER, R. M. BROWN, A. D. DEMEGILLO, AND M. GAULKE. 2006. Gekko ernstkelleri sp. N.—a new gekkonid lizard from Panay Island, Philippines. Salamandra 42:193–206. SILER, C. D., C. W. LINKEM, A. C. DIESMOS, AND A. C. ALCALA. 2007. A new species of Platymantis (Amphibia: Anura: Ranidae) from Panay Island, Philippines. Herpetologica 63:351–364. ZWEIFEL, R. G. 1960. Results of the Gilliard New Britain expedition 3. Notes on the frogs of New Britain. American Museum Novitates 2023:1–27. ZWEIFEL, R. G. 1969. Frogs of the genus Platymantis (Ranidae) in New Guinea, with the description of a new species. American Museum Novitates 2374:1–19. ZWEIFEL, R. G. 1975. Frogs of the genus Platymantis (Ranidae) from New Britain. American Museum Novitates 7582:1–7. .Accepted: 13 February 2009 .Associate Editor: Michael Harvey
APPENDIX I Specimens Examined All specimens examined are from the Philippines. Numbers in parentheses indicate the number of specimens examined for each species. Platymantis banahao.—(6) LUZON ISLAND, QUEZON PROVINCE, Municipality of Tayabas, Baragany Lalo, Mt. Banahao: TNHC 61968–71, PNM 9248–49. Platymantis cagayanensis.— (9) LUZON ISLAND, CAGAYAN PROVINCE, ‘‘Tagat Forest Reserve near Santa Praxedes Town:’’ PNM 7564, 7578, 7496–99, 7506, 7608, 7526. Platymantis cornutus.—(3) LUZON ISLAND, KALINGA PROVINCE, Municipality of Balbalan, Barangay Balbalan: CAS 231498, 231501, CMNH 8128. Platymantis 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: KU 300350, 300352–54; NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Municipality of Valencia, Barangay Bongbong, Camp Lookout, Cuernos de Negros Mt. Range, Mt. Talinis, 500 m elevation: TNHC 61972–87. Platymantis diesmosi.— (10) LUZON ISLAND, ALBAY PROVINCE, Municipality of Tiwi, Barangay Banhaw: PNM 8499 (Holotype), 8500–1 (Paratypes), TNHC 62040–42 (Paratypes), UPLB-MNH 16, 21–23 (Paratypes). Platymantis dorsalis.—(22)
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NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Municipality of Valencia, Barangay Bongbong, Sitio Tagaytay, Mt. Talinis, 1150 m elevation, 9u 159 360 N, 123u 129 1960 E: KU 300356–300377. Platymantis guentheri.—(19) DINAGAT ISLAND, SURIGAO PROVINCE, Municipality of Loreto, Barangay Esperanza: KU 306321–23, 306325; SAMAR ISLAND, EASTERN SAMAR PROVINCE, Municipality of Taft, Barangay San Rafael: KU 309185, 309189, 309191, 309195, 309203, 309217–19, 309221, 309228–30, 309232, 309236, 309238. Platymantis hazelae.—(13) NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Municipality of Valencia, Barangay Bongbong, Sitio Tagaytay, Mt. Talinis, 1150 m elevation: KU 300403–15. Platymantis indeprensus.—(8) LUZON ISLAND, QUEZON PROVINCE, Municipality of Tayabas, Barangay Lalo, Mt. Banahao: TNHC 061956–60; PNM 9257–59. Platymantis insulatus.—(18) SOUTH GIGANTE ISLAND, ILOILO PROVINCE, Municipality of Carles, Barangay Gabi: CAS 117441 (Holotype), 119967–69 (Paratypes); KU 300338–44, 300346, 309088–89; NORTH GIGANTE ISLAND, ILOILO PROVINCE, Municipality of Carles, Barangay Granada: KU 300345, 300347–49. Platymantis isarog.—(7) LUZON ISLAND, CAMARINE SUR PROVINCE, Municipality of Naga City, Barangay Panicuason, Mt. Isarog National Park, Mt. Isarog: TNHC 61961–67. Platymantis lawtoni.—(1) TABLAS ISLAND, ROMBLON PROVINCE, Municipality of San Agustin, Mt. Progreso: CAS 135733. Platymantis levigatus.—(15) SIBUYAN ISLAND, ROMBLON PROVINCE, Municipality of Magdiwang, Barangay Talaba, Mt. Guiting-Guiting Natural Park, 0 m elevation: KU 300416–30. Platymantis luzonensis.—(27) LUZON ISLAND, LAGUNA PROVINCE, Municipality of Los Ban˜os, Barangay Batong Malake, Mt. Makiling: CAS 196364, 196369–70, 200404–08, 210544–45 (Paratypes); CAMARINES SUR PROVINCE, Municipality of Naga City, Mt. Isarog: FMNH 251643–44; TNHC 62004–09, 62012–13, 62020–24; POLILLO ISLAND, QUEZON PROVINCE, Municipality of Polillo, Barangay Pinaglubayan: KU 305541–42. Platymantis mimulus.—(12) LUZON ISLAND, LAGUNA PROVINCE, Municipality of Los Ban˜os, Barangay Batong Malake, Mt. Makiling: TNHC 54930–31; PNM 9260–69. Platymantis montana.—(13) LUZON ISLAND, QUEZON
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PROVINCE, Municipality of Tayabas, Barangay Lalo, 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, Cuernos de Negros Mountain Range, Mt. Talinis, 750 m elevation: KU 300439–45. Platymantis paengi.— (15) PANAY ISLAND, ANTIQUE PROVINCE, Municipality of Pandan, Barangay Duyong: PNM 9239 (Holotype), 9240– 43 (Paratopotypes), KU 300206–13 (Paratopotypes), 300204–05 (Paratypes). Platymantis panayensis.—(2) PANAY ISLAND, AKLAN PROVINCE, Municipality of Nabas: CAS 137641–42. Platymantis pseudodorsalis.—(4) LUZON ISLAND, QUEZON PROVINCE, Municipality of Tayabas, 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, 9528–31. Platymantis rabori.—(12) MINDANAO ISLAND, NEW BATAAN PROVINCE, Mt. Puting Bato: CMNH 2305, 2350; DAVAO CITY PROVINCE, Municipality of Toril, Barangay Baracatan, Sitio Upper Baracatan: CMNH 1462; Municipality of Calinan, Barangay Malagos: PNM 9504–05; SAMAR ISLAND, EASTERN SAMAR PROVINCE, Municipality of Taft, Barangay San Rafael: KU 309121–27. Platymantis sierramadrensis.—(6) LUZON ISLAND, AURORA PROVINCE, Municipality of San Luis; Dipiningan branch of the Kobatangan River drainage; 15u 409 120 N, 121u 209 480 E: CMNH 5678–79, 5904; ISABELA PROVINCE, Municipality ofPalanan, Barangay Didian, Sitio Natap Dukan, Northern Sierra Madre National Park, 16u 579 55.80 N, 122u 249 13.80 E: CAS 204739–41. Platymantis spelaeus.—(7) NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Municipality of Basay, Tiyabanan Barrio: CAS 153477–78, 153482 (Paratypes); NEGROS OCCIDENTAL PROVINCE, Municipality of Cauayan, Sitio Banso, Barangay Camalandaan, 320 m elevation: KU 300435–38. Platymantis subterrestris.—(3) LUZON ISLAND, MOUNTAIN PROVINCE, Mt Data: CAS 204319–204321. Platymantis taylori.—(4) LUZON ISLAND, ISABELA PROVINCE, Municipality of Palanan, Barangay Didian, Sitio Natapdukan: CAS 207443–207446 (Paratypes).
