Herpetologica, 65(1), 2009, 92–104 E 2009 by The Herpetologists’ League, Inc.

A NEW SPECIES OF LIMESTONE-FOREST , GENUS (AMPHIBIA: ANURA: CERATOBATRACHIDAE) FROM EASTERN SAMAR ISLAND, PHILIPPINES

1,3,4 3 2,3 1,3 CAMERON D. SILER ,ANGEL C. ALCALA ,ARVIN C. DIESMOS , AND RAFE M. BROWN 1Natural History Museum and Biodiversity Research Center, Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66045-7561, USA 2Herpetology Section, Zoology Division, Philippine National Museum, Rizal Park, Burgos St., Manila, Philippines 3Angelo King Center for Research and Environmental Management, Silliman University, Dumaguete City, 6200, Philippines

ABSTRACT: A new species of forest frog of the genus Platymantis is described from an elevation of 140 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, salmon- colored 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 ; Platymantis guentheri Species Group; SW Pacific

FROGS of the genus Platymantis have two morphology (W. C. Brown et al., 1997a,c; geographic centers of diversity and endemism. Alcala and W. C. Brown, 1999)—the P. One large group of species is situated in the dorsalis Group, the P. hazelae Group, and (W. C. Brown, 1952, 1997; the P. guentheri Group. In addition to the Menzies, 2007) and nearby Bismarck (Zweifel, morphological variation among Platymantis in 1960, 1975; Foufopoulos and R. M. Brown, the Philippines, the advertisement calls and 2004; R. M. Brown et al., 2006a,b; Foufopou- preferred microhabitats of the species also los and Richards, 2007) and Admiralty archi- vary (R. M. Brown et al., 2002; R. M. Brown pelagos (26 species; Allison, 1996; Richards et and Gonzales, 2007). Surveys have shown that al., 2007). Another large group of species limestone-forest habitats of the Philippines occurs in the Philippines (27 species; Alcala possess highly distinctive species of Platyman- and W. C. Brown, 1998, 1999; W. C. Brown et tis (W. C. Brown and Alcala, 1982; R. M. al., 1997a; R. M. Brown and Gonzales, 2007). Brown and Alcala, 2000; Siler et al., 2007). Outside of these two large archipelagos, one In 1998 the late Walter C. Brown (see species occurs in Palau (Crombie and Pregill, Alcala, 2004) informed RMB about a pair of 1999); seven or eight are known from New highly distinctive Platymantis specimens from Guinea faunal region (Zweifel, 1969; Allison, the Island of Samar that he had examined 1996; Gu¨ nther, 1999, 2005; Menzies, 2007); many years prior. He remarked that the two to four species occur in eastern Indonesia suspected new species was ‘‘a giant form that (Edgar and Lilley, 1993; Menzies, 1982a,b; looked like a cross between Platymantis including undescribed taxa; M. I. Setiadi and speleaus and .’’ Although D. T. Iskandar, personal communication); and Brown could not remember where the spec- two others occur in Fiji (Gorham, 1965; imens were deposited, his verbal description Morrison, 2003). of the suspected new species of giant Platy- Within the Philippines, three species mantis from Samar was quite specific and very groups are recognized on the basis of external detailed. Over the past decade we have been unable to locate the specimens in U.S. collections, but we have anxiously awaited 4 CORRESPONDENCE: e-mail, [email protected] the discovery of the new species that Brown

92 March 2009] HERPETOLOGICA 93

observer bias and other sources of potential error (Hayek et al., 2001; Lee, 1982), all measurements were scored by CDS. Descrip- tions 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), interor- bital distance (IOD), internarial distance (IND), eye diameter (ED), horizontal tym- panic annulus diameter (TAD), eye–tympa- num distance (ETD), head width (HW), upper arm length (UAL), forearm length (FAL), tibia length (TBL), femur length (FEL), tarsus length (TSL), pes length (PL), manus length (ML), Toe IV length (Toe4L), Finger I length (Fin1L), Finger III length (Fin3L), Finger I disc width (Fin1DW), Finger III disc width (Fin3DW), Toe IV disc width (Toe4DW), and widths of penultimate FIG. 1.—Known occurrence of Platymantis bayani in phalanges of third finger (PpFin3) and the eastern Samar Island, Philippines (black dot). The inset shows the location of Samar Island (colored black) within fourth toe (PpToe4). In the description, the Philippines. The three provinces of Samar Island are ranges are followed by mean 6 standard indicated by dashed lines. deviation in parentheses. Specimens were deposited at the Philippine National Museum predicted, ‘‘would eventually turn up on (PNM) and Kansas University (KU). For this Samar.’’ description, we follow Kraus and Allison During the month of October 2007, we (2007) in maintaining the masculine forma- conducted herpetological field surveys in the tions of species names within the genus lower-elevation forest of the Municipality of Platymantis. Taft in eastern Samar Island (Fig. 1). Twenty- one individuals of an undescribed species of RESULTS Platymantis that match Walter Brown’s de- Platymantis bayani sp. nov. scription were collected at an elevation of (Figs. 2–4) 140 m. Despite its striking tree frog appear- Holotype.—PNM 9501 (field no. CDS ance, the new species is found in terrestrial 2989, formerly KU 309260; Fig. 2), an adult microhabitats in karst forest, an ecomorpho- male collected at 20:00 hr on 22 October 2007 logical combination only shared with P. at 140 m elevation in Taft Forest (11u 489 insulatus (R. M. Brown and Alcala, 2000). 9.180 N, 125u 179 33.9360 E; WGS-84) Herein, we describe the new species and Barangay San Rafael, Municipality of Taft, report on its natural history, ecology, and Eastern Samar Province, Samar Island, Phi- habitat. lippines, by Cameron D. Siler and Jason B. Fernandez. MATERIALS AND METHODS Paratopotypes.—KU 309252–54, 309256, We recorded morphometric data from 309258–59, 309261, 309263–64, 309266–67, alcohol-preserved specimens that were fixed and 309269, PNM 9515–22, eleven males and in 10% formalin (Appendix 1). Sex was nine females, collected between 14 and 23 determined by gonadal inspection, and mea- October 2007. surements were taken with digital calipers to Diagnosis.—Platymantis bayani can be the nearest 0.1 mm. To minimize inter- distinguished from other Philippine congeners 94 HERPETOLOGICA [Vol. 65, No. 1

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; supra- tympanic fold moderately protuberant; tym- panum completely visible; dorsolateral stripes absent; and a preference for terrestrial microhabitat in karst forest. Mensural and diagnostic differences are provided in Ta- bles 1 and 2. Comparisons.—The new species is assign- FIG. 2.—Head of male holotype of Platymantis bayani (PNM 9501) in oblique view. Scale bar 5 5 mm. able to the P. guentheri Group (P. banahao, P. cornutus, P. diesmosi, P. guentheri, P. insula- tus, P. luzonensis, P. negrosensis, and P. by the following combination of characters: rabori) based on the following combination body size 34.2–39.1 mm SVL for males, 44.3– of characters: digital discs greatly expanded 49.8 mm SVL for females; digital disc of and larger in fingers than in toes, first finger Finger III 2.7–4.03 width of penultimate only slightly shorter than second, proximal phalanx in males and 2.8–4.53 in females; portions of digits with narrow dermal folds in digital disc of Toe IV 2.1–3.33 width of cross section, and subarticular tubercles large penultimate phalanx in males and 2.9–3.73 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. March 2009] HERPETOLOGICA 95

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. 6HREOOIA[ HERPETOLOGICA 96

TABLE 1.—Summary of qualitative diagnostic characters (present 5 +, absent 52)inPlatymantis 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.

bayani sp. nov. insulatus negrosensis luzonensis rabori guentheri banahao cornutus diesmosi (9m, 9f) (8m, 6f) (3m, 2f) (9m, 2f) (2m, 6f) (11m, 8f) (4m) (3m) (8m, 2f) Range Samar Gigantes Negros and Luzon Mindanao Mindanao SE Luzon N. Luzon SE Luzon Island Islands Panay Islands PAIC PAIC PAIC Island Island Island SVL (m) 34.2–39.1 34.3–38.0 28.3–35.7 24.1–32.5 30.4–33.5 26.5–29.8 32.4–36.3 26.3–30.4 28.5–39.2 (37.1 6 1.8) (36.7 6 1.3) (31.1 6 4.0) (29.7 6 2.4) (32.0 6 2.2) (28.6 6 1.1) (33.9 6 1.7) (27.8 6 2.2) (35.3 6 3.6) SVL (f) 44.3–49.8 40.2–42.9 41.7–42.2 32.6–33.8 36.4–44.4 32.3–37.3 N/A N/A 49.0–52.6 (46.4 6 2.1) (41.8 6 1.0) (42.0 6 0.3) (33.2 6 0.8) (41.5 6 3.6) (34.6 6 2.0) (50.8 6 2.6) ED (m) 4.8–6.1 3.7–4.9 3.1–4.1 2.7–3.9 3.8–3.9 2.8–3.6 3.6–4.1 3.0–4.2 3.7–5.4 (5.5 6 0.5) (4.2 6 0.4) (3.7 6 0.6) (3.4 6 0.4) (3.9 6 0.1) (3.3 6 0.3) (3.8 6 0.2) (3.5 6 0.6) (4.7 6 0.5) ED (f) 5.4–6.8 5.1–5.7 4.4–4.7 3.6–3.9 4.0–5.7 3.7–4.5 N/A N/A 5.9–6.1 (6.1 6 0.4) (5.3 6 0.2) (4.6 6 0.2) (3.7 6 0.2) (4.9 6 0.7) (3.9 6 0.3) (6.0 6 0.1) Fin3DW/PpFin3 2.7–4.0 2.7–3.1 2.6–2.8 2.2–3.3 2.8–3.5 2.4–5.2 2.8–3.2 2.5–2.9 1.5–2.1 (m) (3.3 6 0.4) (2.9 6 0.1) (2.7 6 0.1) (2.8 6 0.4) (3.2 6 0.5) (3.2 6 0.8) (3.0 6 0.2) (2.7 6 0.2) (1.7 6 0.2) Fin3DW/PpFin3 2.8–4.5 2.3–2.8 3.5–4.7 3.0–3.2 3.3–4.2 2.6–5.7 N/A N/A 1.6–1.7 (f) (3.9 6 0.5) (2.7 6 0.2) (4.1 6 0.8) (3.1 6 0.2) (3.6 6 0.3) (4.0 6 1.2) (1.6 6 0.1) Toe4DW/PpToe4 2.1–3.3 1.9–3.4 1.8–1.9 1.6–2.2 2.0–2.7 1.5–3.5 1.9–2.3 1.6–2.0 1.2–2.2 (m) (2.8 6 0.4) (2.6 6 0.4) (1.8 6 0.1) (1.9 6 0.2) (2.3 6 0.5) (2.4 6 0.6) (2.0 6 0.2) (1.7 6 0.2) (1.7 6 0.4) Toe4DW/PpToe4 2.9–3.7 2.0–2.6 1.8–1.8 1.8–1.9 1.6–2.5 2.3–4.3 N/A N/A 1.7–2.1 (f) (3.1 6 0.3) (2.3 6 0.3) (1.8 6 0.0) (1.8 6 0.1) (2.3 6 0.3) (3.2 6 0.7) (1.9 6 0.3) Fin3DW/Toe4DW 1.1–1.5 1.1–1.5 1.8–2.0 1.6–2.0 1.3–1.6 1.2–1.7 1.8–1.9 1.9–2.1 0.9–1.4 (m) (1.3 6 0.1) (1.3 6 0.1) (1.9 6 0.1) (1.7 6 0.1) (1.4 6 0.2) (1.4 6 0.1) (1.8 6 0.0) (2.0 6 0.1) (1.1 6 0.2) Fin3DW/Toe4DW 1.2–1.5 1.2–1.6 1.7–2.0 2.0–2.1 1.4–1.9 1.1–1.7 N/A N/A 0.8–0.8 (f) (1.3 6 0.1) (1.4 6 0.1) (1.9 6 0.3) (2.1 6 0.1) (1.6 6 0.2) (1.4 6 0.2) (0.8 6 0.0) Greatly expanded digital disks ++ ++++, moderate ++2 Dorsal skin rugosity 2 to granular 2 ++++2 to granular 22to granular Dorsal dermal tubercles +, sparse 2 +++, dense ++22 Tubercle color Salmon N/A Cream Dominant Dominant Dominant Dominant N/A N/A dorsal color dorsal color dorsal color dorsal color 22 ++++222 Dorsal folds/ridges 1 No. 65, Vol. Tympanum hidden No No Posterodorsal Posteroventral Posterior Posteroventral Posterodorsal No Posterodorsal edge edge edge edge edge edge Supratympanic Smooth, less Smooth, not Rugose, prominent Rugose, Rugose, Smooth, less Smooth, less Smooth, not Smooth, less fold prominent prominent prominent prominent prominent prominent prominent prominent March 2009] HERPETOLOGICA 97

iff From all members of the P. dorsalis Group (P. cagayensis, P. corrugatus, P. dorsalis, P. 2 edges ,well

defined indeprensus, P. levigatus, P. mimulus, P. (8m, 2f) diesmosi + naomiae, P. paengi, P. pseudodorsalis, P. pyg- maeus, P. spelaeus, and P. taylori), Platyman- tis bayani is distinguished by the presence (vs. absence) of the following features: greatly

+ expanded digital discs (Fig. 3); slightly larger (3m) defined , poorly cornutus

+ 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 2

faint color in P. dorsalis, P. paengi, P. spelaeus,and or (4m) banahao

+ 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 22 further distinguished from all members of the or , well defined + (11m, 8f) + guentheri 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 sub- 2 strates in limestone forest vs. leaf litter of or , well defined rabori (2m, 6f) + + forest floor and/or herb-layer vegetation in P. cagayensis, P. corrugatus, P. dorsalis, P. inde- prensus, P. mimulus, P. naomiae, P. pseudo- dorsalis, P. pygmaeus and P. taylori, or river 1.—Continued. 2

or banks and rocks in P. levigatus). From P. ,well defined (9m, 2f) + ABLE luzonensis cagayensis, P. corrugatus, P. dorsalis, P. inde- T prensus, P. mimulus, P. naomiae, P. pygmaeus and P. taylori, the new species is distinguished by the absence (vs. presence) of dorsal folds

++ and ridges. 2222222 ,well defined (3m, 2f) + The new species is distinguished from all P. negrosensis Arboreal Arboreal Arboreal Shrub/Arboreal Arboreal Arboreal Ground; cl hazelae Group species (P. hazelae, P. isarog, P. lawtoni, P. montanus, P. panayensis, P. poli- llensis, P. sierramadrensis, P. subterrestris)by its larger body size; the first finger much , poorly habitat defined 2 shorter than the second; narrow dermal (8m, 6f) insulatus

or phalanges on the digits; finger discs only Ground; karst + slightly larger than toe disks; protuberant (vs. flattened) subarticular tubercles, and a pre- ferred microhabitat of terrestrial substrates in ++ 22 limestone forest (vs. arboreal habitat, and/or habitat (9m, 9f) shrub-layer vegetation). bayani sp. nov. , well defined Ground; karst + 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 stripes preference species of the P. guentheri Group, except for Dorsolateral Spotted flanks Limb bands Microhabitat P. diesmosi, by its larger body size and greater 98 HERPETOLOGICA [Vol. 65, No. 1

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).

insulatus bayani sp. nov. (9m, 9f) (8m, 6f) spelaeus (4m, 3f) paengi (15m) SVL (m) 34.2–39.1 34.3–38.0 37.1–44.3 27.7–34.3 SVL (f) 44.3–49.8 40.2–42.9 53.5–57.5 N/A Range Samar Island Gigantes Islands SW Negros Island NW Panay Island Species Group guentheri guentheri dorsalis dorsalis Digital disks Widely expanded Widely expanded Non-expanded Non-expanded Dorsal skin rugosity 2 to granular 2 +, 2 + Dorsal dermal tubercles +, sparse 2 +, small + Flanks Spotted Spotted Immaculate Spotted eye diameter. The new species differs from P. eyes large and protruding laterally, ED 15.6% negrosensis, P. luzonensis, P. rabori, and P. SVL, 41.3% HL; lips slightly flared and guentheri by lacking rugose dorsal skin, dorsal swollen (Fig. 2); interorbital region flat; ED folds and ridges, and dorsolateral stripes. It is 137.0% SNL, 1.73 END; pupil horizontally distinguished from P. diesmosi, P. negrosensis, elliptical; canthus rostralis laterally concave in P. luzonensis, P. rabori, P. guentheri, and P. dorsal aspect; loreal region greatly concave, banahao by having a completely visible sloping ventrolaterally to labium; nostrils tympanum, dark brown to black spots on the oriented posterolaterally, protruding laterally flanks, sparsely distributed dorsal dermal to sharp points from snout tip; END 2.33 tubercles that are salmon-colored, and a SNL; internarial region flat; tympanic annulus preference for terrestrial microhabitats in distinct, its diameter 0.53 ED; supratympanic limestone forest. Platymantis bayani is further fold moderately protuberant, extending from distinguished from P. negrosensis, P. luzonen- posterior corner of eye across dorsal margin of sis, and P. rabori by having a smooth and less tympanum to supra-axillary region. prominent supratympanic fold. From P. cor- Tongue elongate, wider posteriorly, with nutus, P. negrosensis, P. luzonensis,andP. deep posterior notch and narrow anterior banahao, it differs in having slightly larger attachment; choanae situated at anterolateral finger discs than toe discs. The new species edge of palate, oval, widely separated by differs from P. diesmosi in having greatly distance three to four times greater than expanded digital discs, and from P. cornutus, diameter of single choana; dentigerous pro- by having dark brown to black spots on the cess of vomer distinct, with two teeth located flanks, sparsely distributed, salmon-colored, on anterior 1/3 of process; dentigerous process dorsal dermal tubercles, a slightly prominent angled slightly anterolaterally, approximately supratympanic fold, well-defined limb bands, at 10–20u incline with closest (posterior) the absence of dorsolateral stripes, and a points separated by distance equal to 2.03 preference for terrestrial microhabitats in diameter of single choana, their most distant limestone forest. From P. insulatus, the new (anterior) ends separated by distance equal to species is distinguished by the presence of 3.03 diameter of single choana; short vocal dorsal dermal tubercles and a slightly prom- slits at posteroventral margin of mouth. inent supratympanic fold. Finally, P. bayani Skin of dorsal surfaces of trunk and head differs from P. insulatus and P. banahao in coarsely granular, distinctly shagreen; irregular, having a distinct, transverse limb-banding moderately protuberant tubercles sparsely dis- pattern. tributed across dorsal surfaces of body and limbs; Description of holotype.—A mature male; ventral surfaces of head smooth; lateral and habitus robust; head broader than body, HL ventral surfaces of limbs finely granular with 37.7% SVL, 91.2% HW; snout protruding dorsal surfaces coarsely granular; flanks shagreen. moderately beyond lower jaw, snout tip comes Manus length 58.0% pes length; tibia length to sharp point in dorsal and lateral aspects; 108.8% pes length; tibia length 54.6% SVL; March 2009] HERPETOLOGICA 99 fingers slender, long; terminal discs greatly dark brown and black; lips lighter gray with expanded (Fig. 3A) with pronounced distal, faint vertical dark brown labial bars and circummarginal groove; dorsal surfaces of speckles, bordered anteriorly by lighter brown terminal phalanges with small, cutaneous lower edge of canthus and subocular regions; supra-articular flaps between ultimate and transverse, dark brown bars on limbs similar penultimate phalanges; relative lengths of in color to dorsum, coalescing into mottled fingers I 5 II , IV , III; subarticular patterns on elbows and knees; four bars on tubercles prominent, convex, granular in forearms, four on thigh, four on tibia, three on texture; one subarticular tubercle below tarsus; spotted flanks, ground color lighter Digits I and II, two tubercles under Digits grayish tan with dark brown blotches and III and IV; supernumerary tubercles distinct, spots, blending into the darker dorsal pattern round, prominent; present at bases of Digits (above) and the lighter ventral pattern (be- I–IV; inner metacarpal tubercle enlarged, low); dorsal surfaces of manus, pes, and digits elongate, crescent shaped; outer metacarpal light grayish tan with dark brown blotches tubercle partially divided into oval, slightly above each joint. raised preaxial tubercle of moderate size, and Throat marbled light cream to tan with round postaxial tubercle equal in size to speckled green-brown blotches; periphery of preaxial metacarpal tubercle; nuptial pads throat and lower lips lighter with anterior absent; forearms slender, not hypertrophied edge of lower lip possessing outline of light in males. cream spots; chest light cream with less dense Tarsus smooth, lacking folds, flaps, or speckled green-brown blotches; lighter ventral tubercles; terminal discs of toes greatly coloration limited to small area on ventral expanded, with circummarginal grooves; su- surface of forelimb near limb insertion; pra-articular cutaneous flaps above ultimate- ventral surface of upper- and forearms darker penultimate phalangeal articulations; plantar cream with green-brown blotches; venter light surface of foot smooth, with well-developed, cream with nearly indistinct light green-brown prominently rounded subarticular tubercles blotches that decrease in size posteriorly until (Fig. 3B); supernumerary tubercles absent; blending with uniformly light cream groin; relative lengths of toes I , II , V , III , ventral surfaces of thigh uniformly mottled IV; outer metatarsal tubercle small, round; cream and green-brown; ventral surface of inner metatarsal tubercle prominent, with tibia and tarsus mottled green-brown, similar sharp inner edge, length equal to distance to forearms; palmar and plantar surfaces of from base of Toe I to first subarticular hands and feet dark green-brown with light tubercle; toes with vestige of basal interpha- cream subarticular tubercles; iris pale gray langeal web, nearly extending to the level of above and dark gray below pupil. the proximal subarticular tubercles between Coloration of holotype in life.—Dorsum Toes III and IV. and head mottled light green, dark green, Coloration of holotype in preservative.— and black (Fig. 4); interorbital bar distinct, Dominant dorsal color pattern on head and black, and bordered anteriorly by lighter, body mottled with gray and silver blotches and green-brown border; dorsal surfaces of limbs spots interspersed throughout a dominant with alternating dark brown and green trans- background of black and dark brown; dorsal verse bars; dorsal surface of body with low, surfaces of limbs banded with alternating salmon-colored tubercles; series of dark green-brown and dark brown transverse bars; brown to black blotches and spots across interorbital bar slightly darker than dominant flanks; pupil bordered by bright gold ciliary dorsal color pattern; snout region slightly ring of iris, blending into the predominant darker than dorsum, and nearly uniform mottled golden-brown and dark brown color black; wide black bar extending from snout pattern of iris. tip, across canthus rostralis to anterior edge of Measurements of holotype (mm).—SVL eye; a dark blotch covering anteroventral edge 39.1; HL 14.7; HW 16.2; SNL 4.4; IND 3.6; of tympanum, ventral two thirds of tympanum IOD 3.2; ED 6.1; END 3.5; TAD 2.8; ETD lighter gray; temporal region dorsally mottled 1.0; UAL 7.4; FAL 9.0; FEL 20.1; TBL 21.3; 100 HERPETOLOGICA [Vol. 65, No. 1

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 Male n 5 9 SVL 44.3–49.8 (46.4 6 2.1) 34.2–39.1 (37.1 6 1.8) HL 16.2–18.8 (17.3 6 0.8) 14.1–15.5 (14.8 6 0.5) END 3.3–4.8 (4.1 6 0.5) 3.2–4.4 (3.9 6 0.4) SNL 3.8–6.1 (5.3 6 0.8) 4.0–5.9 (5.0 6 0.8) IOD 4.0–4.9 (4.3 6 0.3) 3.0–4.2 (3.5 6 0.3) IND 3.6–4.3 (3.9 6 0.2) 2.9–3.6 (3.2 6 0.2) ED 5.4–6.8 (6.1 6 0.4) 4.8–6.1 (5.5 6 0.5) TAD 2.3–3.3 (2.8 6 0.3) 1.9–2.8 (2.4 6 0.3) ETD 1.1–1.7 (1.4 6 0.2) 1.0–1.6 (1.3 6 0.2) HW 18.4–20.6 (19.7 6 0.7) 15.1–17.0 (15.9 6 0.6) UA 8.2–9.9 (9.1 6 0.6) 7.3–8.1 (7.6 6 0.3) FAL 12.0–13.1 (12.6 6 0.4) 9.8–10.6 (10.3 6 0.2) FL 23.7–26.2 (24.9 6 0.9) 19.6–20.7 (20.2 6 0.4) TBL 21.0–26.9 (25.8 6 1.8) 21.2–22.8 (21.7 6 0.5) TSL 14.7–15.9 (15.1 6 0.4) 12.0–13.2 (12.6 6 0.4) PL 22.9–25.3 (23.9 6 0.9) 18.7–20.3 (19.7 6 0.6) ML 13.5–14.8 (14.2 6 0.3) 11.2–12.3 (11.7 6 0.4) Toe4L 21.8–23.9 (22.8 6 0.6) 18.1–20.0 (18.8 6 0.6) Fin1L 7.2–8.4 (7.9 6 0.4) 6.1–7.1 (6.4 6 0.4) Fin3L 12.7–13.9 (13.2 6 0.4) 10.3–11.5 (10.8 6 0.4) Fin1DW 1.6–2.0 (1.8 6 0.1) 1.3–1.6 (1.4 6 0.1) Fin3DW 2.3–2.9 (2.7 6 0.2) 1.8–2.5 (2.1 6 0.2) Toe4DW 1.7–2.3 (2.0 6 0.2) 1.5–1.8 (1.6 6 0.1) PpFin3 0.6–0.8 (0.7 6 0.1) 0.5–0.7 (0.7 6 0.1) PpToe4 0.6–0.7 (0.6 6 0.1) 0.5–0.7 (0.6 6 0.1)

TSL 12.8; ML 11.4; PL 19.6; Fin1L 6.1; new species for the late Walter C. Brown in Fin3L 10.3; Toe4L 18.2; Fin3DW 1.8; recognition of his numerous contributions to Toe4DW 1.5; PpFin3 0.7; PpToe4 0.7. Philippine herpetology and in acknowledg- Variation.—Summaries of univariate mor- ment of his lifelong commitment to the study phological variation in the series are presented of biodiversity in the SW Pacific. Suggested in Table 3. Dominant color pattern consis- common name: Walter’s Limestone Forest tently mottled in all observed specimens; Frog. however, the degree of contrast between the Ecology and natural history.—Platymantis dominant ground color and dark blotches bayani occurs in primary- and secondary- varies among individuals, with two patterns of growth forest on karst substrate. The adver- contrast observed. The first pattern consists of tisement call of the species is unknown. Both dark brown to black blotches intermixed with males and females were found perched on top a light cream to green ground color (KU of limestone rocks within large rock forma- 309253–54, 309256, 309258–59, 309261, and tions in the forest; they never were observed 309264). The second pattern was consists of perching in trees or shrubs as do most species dark green blotches intermixed with the same in the P. guentheri Group (R. M. Brown and light cream to green ground color (KU Gonzales, 2007). 309266, 309269, 309252, 309263, and Platymantis bayani was well camouflaged 309267). against the limestone (much like P. insulatus, Distribution.—Platymantis bayani is known P. speleaus, and P. paengi; Fig. 4), and only from the type locality (Fig. 1). collectors found it difficult to locate and Etymology.—The name of the new species capture individuals against the limestone is derived from the Tagalog (Filipino) term substrate. When disturbed, the new species ‘‘bayani,’’ meaning ‘‘highly respected,’’ ‘‘hero- immediately jumped back into the rocks and ic,’’ or ‘‘hero.’’ We are pleased to name this escaped into limestone crevices. No egg March 2009] HERPETOLOGICA 101 masses or juveniles were found; other species known karst-forest frogs found in the Visayan of Platymantis have direct development (W. (Central) Philippine Islands. All four species C. Brown and Alcala, 1982a,b). Platymantis seem to be obligate inhabitants of limestone spelaeus has been observed by one of us forests. Although potentially restricted in (CDS) to deposit direct-developing eggs in geographic ranges, the species are quite limestone crevices and the same is suspected common in the rainy season, but can seem for P. insulatus (R. M. Brown and Alcala, to be rare in times of high temperature and 2000). We assume that this species also breeds aridity when we assume they retreat into in limestone crevices and small caves (R. M. subterranean limestone caves and crevices (R. Brown and Alcala, 2000; R. M. Brown et al., M. Brown and Alcala, 2000; R. M. Brown et 2003). al., 2003). Sympatric anuran species include Kalo- The recent discoveries of new species of phrynus pleurostigma, Limnonectes magnus, Philippine frogs and gekkos with preferences Megophrys stejnegeri, Platymantis cf. dorsalis, for limestone forests (Siler et al., 2007; P. cf. corrugatus, P. rabori, P. guentheri, Roesler et al., 2006) highlight the degree to Polypedates leucomystax, Philautus leitensis, which this imperiled habitat type (Clements et Rhacophorus bimaculatus, R. appendiculatus, al., 2006) is poorly known. Future survey R. pardalis, Oreophryne sp., Rana grandocula, efforts should make a point to target limestone R. albotuberculata, and Occidozyga laevis. forest communities. Our general impression is that these habitat types have been overlooked DISCUSSION in the past because surrounding forests are With the new species described here, a total dry, stunted, difficult to sample, and viewed of 28 Philippine species of Platymantis are by field biologists as non-productive. now recognized. The species total for the The heavily mottled, predominant color country has increased dramatically over the pattern of Platymantis bayani resembles P. last several decades; seven species were spelaeus, P. insulatus, and P. paengi. The recognized between 1950 and 1970 (Inger, conspicuous similarity of color patterns among 1954; W. C. Brown and Alcala, 1970a,b)and these taxa suggests to us the possibility of 12 by the late 1980’s (W. C. Brown and Alcala, convergent selection on color pattern in 1982a). An additional dozen species were limestone habitats. It is unclear as to which described in the late 1990’s (W. C. Brown et species P. bayani is most closely related, but al., 1997a,b,c, 1999), several more species phenotypic similarity with P. insulatus is have been described recently (R. M. Brown striking. and Gonzales, 2007; Siler et al., 2007), and The new species was quite common at the numerous unnamed species await description type locality, and it may posses a wider (Brown, 2004; Brown et al., 2008). geographical distribution, possibly throughout Within the Philippine radiation, high levels the northern islands of the Mindanao Faunal of cryptic diversity (Alcala et al., 1998; W. C. Region (Samar, Leyte, Biliran, Mindanao, Brown et al., 1997b, 1999) has only recently Dinagat, Siargao, and smaller associated been uncovered (Brown, 2004; Brown et al., islands). It is possible that a primary limiting 2008) due to attention on advertisement-call factor to the range of P. bayani is the presence diversity, microhabitat preferences, and gene of relatively undisturbed limestone formations sequence data and forest cover. We note that both P. Platymantis bayani is the fourth Philippine speleaus and P. insulatus persist quite well, limestone frog to be described, the third despite the near complete removal of forest on terrestrial member of the P. guentheri Group the Gigante Island group (P. insulatus) and (the others being P. insulatus and P. diesmosi), southern Negros Island (P. spelaeus). Inten- and the only limestone species known to occur sive survey efforts will be required to fully in the Mindanao Faunal Region. Platymantis document the distribution of the new species spelaeus from SW Negros Island, P. insulatus and arrive at a reasonable assessment of its from the Gigantes Islands, and P. paengi from conservation status. At present, we consider NW Panay Island are the three previously the status of the new species to be ‘‘data 102 HERPETOLOGICA [Vol. 65, No. 1 deficient’’ pending the collection of much Signals of Southeast Asian Forest Frogs (Subfamily needed data on its distribution and habitat Platymantinae). Doctoral dissertation, University of Texas at Austin, Austin, Texas, U.S.A. requirements. BROWN, R. M., AND A. C. ALCALA. 2000. Geckos, cave Acknowledgments.—We thank the Protected Areas and frogs, and small land-bridge islands in the Visayan sea. Wildlife Bureau (PAWB) of the Philippine Department of Haring Ibon 2:19–22. Environment and Natural Resources (DENR) for facili- BROWN, R. M., AND J. C. GONZALES. 2007. A new forest tating collecting and export permits necessary for this and frog of the genus Platymantis (Amphibia; Anura; related studies, wherein we are particularly grateful to M. Ranidae) from the Bicol Peninsula of Luzon Island, Lim, C. Custodio, and A. Tagtag. We also thank the Arnie Philippines. Copeia 2007:251–266. Viojan and the Regional Office of the Protected Areas, BROWN, R. M., A. DIESMOS, AND A. C. ALCALA. 2002. The Wildlife, and Coastal Zones Management Services state of Philippine herpetology and the challenges for (PAWCZMS), and Director Manolito D. Ragub, Eirish the next decade. Haring Ibon 42:18–87. Mate, and the headquarters of the Samar Island Natural BROWN, R. M., A. C. DIESMOS, AND A. C. ALCALA. 2008. Parks (SINP), whose logistical support and guidance were Philippine biodiversity is increasing in leaps instrumental throughout our visit to Samar. Jason and bounds. Pp. 82–83. In S. N. Stuart, M. Hoffmann, Fernandez assisted in the field. Financial support for J. S. Chanson, N. A. Cox, R. Berridge, P. Ramani, and fieldwork was provided by a Panorama Fund grant to B. E. Young (Eds.). Threatened of the CDS from The University of Kansas Natural History World. Lynx Ediciones, Barcelona, Spain; IUCN—The Museum and Biodiversity Institute, a Madison and Lila World Conservation Union, Gland, Switzerland; and Self Fellowship to CDS from the University of Kansas, Conservation International, Arlington, Virginia, U.S.A. NSF EF-0334952 to L. Trueb and R. Brown and DEB BROWN, R. M., C. N. DOLINO,E.ALCALA,A.C.DIESMOS, 0743491 to RMB. For the loans of specimens (museum AND A. C. ALCALA. 2003. The advertisement calls of two abbreviations follow Leviton et al., 1985), we thank J. endangered species of endemic Philippine frogs: Vindum and A. Leviton (CAS), R. Sison and A. C. Platymantis spelaeus and P. insulatus (Anura; Ranidae). Diesmos (PNM), J. Ferner (CMNH), A. Resetar and H. Haring Ibon 43:91–109. Voris (FMNH), R. Crombie (USNM), and T. LaDuc BROWN, R. M., J. FOUFOPOULOS, AND S. J. RICHARDS. 2006. (TNHC). Critical reviews of the manuscript were New species of Platymantis (Amphibia; Anura: Rani- provided by L. Trueb and J. Esselstyn. We thank Michael dae) from New Britain and redescription of the poorly Garfield for illustrating the holotype for Figure 2. CDS known Platymantis nexipus. Copeia 2006:674–695. thanks the Philippine-American Education Foundation, BROWN, R. M., S. J. RICHARDS,J.SUKUMARAN, AND J. A.C. Alcala and family, and M. Diesmos for their FOUFOPOULOS. 2006. A new morphologically cryptic continued support, as well as CAS’s Stearns Fellowship species of forest frog (genus Platymantis) from New for funding a recent visit to examine comparative material. Britain Island, Bismarck Archipelago. Zootaxa Finally, we owe a debt of gratitude to the late Walter C. 1334:45–68. Brown who alerted RMB to the presence of this new BROWN, W. C. 1952. The amphibians of the Solomon species on Samar more than 10 yr before we were able to Islands. Bulletin of the Museum of Comparative find it. Once again, Walter was right. Zoology, Harvard 107:1–64. BROWN, W. C. 1997. 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Banahao: TNHC northern Papua Province, Indonesia (Amphibia: Anura: 61968–71, PNM 9248–49. Platymantis cagayanensis.— Ranidae). Zoologische Abhandlungen 55:85–94. (9) LUZON ISLAND, CAGAYAN PROVINCE, ‘‘Tagat Forest HAYEK, L.-A. C., W. R. HEYER, AND C. GASCON. 2001. Frog Reserve near Santa Praxedes Town:’’ PNM 7564, 7578, morphometrics: a cautionary tale. Alytes 18:153–177. 7496–99, 7506, 7608, 7526. Platymantis cornutus.—(3) NGER I , R. F. 1954. Systematics and zoogeography of LUZON ISLAND, KALINGA PROVINCE, Municipality of Philippine Amphibia. Fieldiana: Zoologica 33:182–531. Balbalan, Barangay Balbalan: CAS 231498, 231501, KRAUS, F., AND A. ALLISON. 2007. Two new species of CMNH 8128. .—(22) CAMI- Platymantis (Anura: Ranidae) from New Britain. GUIN ISLAND, CAMIGUIN PROVINCE, Municipality of Zootaxa 1485:13–32. Guinsiliban, Barangay Cabuan: KU 300351, 300355; LEVITON, A. E., R. H. GIBBS,JR., E. HEAL, AND C. E. POLILLO ISLAND, QUEZON PROVINCE, Municipality of DAWSON. 1985. Standards in herpetology and ichthyol- Polillo, Barangay Pinaglubayan, 62 m elevation, 14u 459 ogy: Part I. Standard symbolic codes for institutional 9.30 N, 121u 589 5.520 E: KU 300350, 300352–54; resource collections in herpetology and ichthyology. NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Munic- Copeia 1985:802–821. ipality of Valencia, Barangay Bongbong, Camp Lookout, LEE, J. C. 1982. Accuracy and precision in anuran Cuernos de Negros Mt. Range, Mt. Talinis, 500 m morphometrics: artifacts of preservation. Systematic elevation: TNHC 61972–87. Platymantis diesmosi.— Zoology 31:266–281. (10) LUZON ISLAND, ALBAY PROVINCE, Municipality of MENZIES, J. I. 1982a. The voices of some male Platymantis Tiwi, Barangay Banhaw: PNM 8499 (Holotype), 8500–1 species of the New Guinea region. British Journal of (Paratypes), TNHC 62040–42 (Paratypes), UPLB-MNH Herpetology 6:1–5. 16, 21–23 (Paratypes). Platymantis dorsalis.—(22) 104 HERPETOLOGICA [Vol. 65, No. 1

NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Munic- PROVINCE, Municipality of Tayabas, Barangay Lalo, Mt. ipality of Valencia, Barangay Bongbong, Sitio Tagaytay, Banahao: TNHC 62149–58; CAS 200998–1000. Platy- Mt. Talinis, 1150 m elevation, 9u 159 360 N, 123u 129 1960 mantis naomiae.—(3) LUZON ISLAND, QUEZON PROV- E: KU 300356–300377. Platymantis guentheri.—(19) INCE, Municipality of Tayabas, Baragany Lalo, Mt. DINAGAT ISLAND, SURIGAO PROVINCE, Municipality of Banahao: TNHC 62169–71. Platymantis negrosen- Loreto, Barangay Esperanza: KU 306321–23, 306325; sis.—(7) NEGROS ISLAND, NEGROS ORIENTAL PROV- SAMAR ISLAND, EASTERN SAMAR PROVINCE, Municipal- INCE, Cuernos de Negros Mountain Range, Mt. Talinis, ity of Taft, Barangay San Rafael: KU 309185, 309189, 750 m elevation: KU 300439–45. Platymantis paengi.— 309191, 309195, 309203, 309217–19, 309221, 309228–30, (15) PANAY ISLAND, ANTIQUE PROVINCE, Municipality of 309232, 309236, 309238. Platymantis hazelae.—(13) Pandan, Barangay Duyong: PNM 9239 (Holotype), 9240– NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Munic- 43 (Paratopotypes), KU 300206–13 (Paratopotypes), ipality of Valencia, Barangay Bongbong, Sitio Tagaytay, 300204–05 (Paratypes). Platymantis panayensis.—(2) Mt. Talinis, 1150 m elevation: KU 300403–15. Platy- PANAY ISLAND, AKLAN PROVINCE, Municipality of mantis indeprensus.—(8) LUZON ISLAND, QUEZON Nabas: CAS 137641–42. Platymantis pseudodorsa- PROVINCE, Municipality of Tayabas, Barangay Lalo, Mt. lis.—(4) LUZON ISLAND, QUEZON PROVINCE, Munici- Banahao: TNHC 061956–60; PNM 9257–59. Platyman- pality of Tayabas, Baragany Lalo, Mt. Banahao: KU tis insulatus.—(18) SOUTH GIGANTE ISLAND, 207455–57, 207459 (Paratypes). Platymantis pyg- ILOILO PROVINCE, Municipality of Carles, Barangay Gabi: maeus.—(5) LUZON ISLAND, CAGAYAN PROVINCE, CAS 117441 (Holotype), 119967–69 (Paratypes); KU Municipality of Calveria, Barangay Mabnang, Mabnang 300338–44, 300346, 309088–89; NORTH GIGANTE Falls: PNM 7523, 9528–31. Platymantis rabori.—(12) ISLAND, ILOILO PROVINCE, Municipality of Carles, MINDANAO ISLAND, NEW BATAAN PROVINCE, Mt. Barangay Granada: KU 300345, 300347–49. Platymantis Puting Bato: CMNH 2305, 2350; DAVAO CITY PROVINCE, isarog.—(7) LUZON ISLAND, CAMARINE SUR PROVINCE, Municipality of Toril, Barangay Baracatan, Sitio Upper Municipality of Naga City, Barangay Panicuason, Mt. Baracatan: CMNH 1462; Municipality of Calinan,Barangay Isarog National Park, Mt. Isarog: TNHC 61961–67. Malagos: PNM 9504–05; SAMAR ISLAND, EASTERN SAMAR Platymantis lawtoni.—(1) TABLAS ISLAND, ROMBLON PROVINCE, Municipality of Taft, Barangay San Rafael: KU PROVINCE, Municipality of San Agustin, Mt. Progreso: 309121–27. Platymantis sierramadrensis.—(6) LUZON CAS 135733. Platymantis levigatus.—(15) SIBUYAN ISLAND, AURORA PROVINCE, Municipality of San Luis; ISLAND, ROMBLON PROVINCE, Municipality of Magdi- Dipiningan branch of the Kobatangan River drainage; 15u wang, Barangay Talaba, Mt. Guiting-Guiting Natural 409 120 N, 121u 209 480 E: CMNH 5678–79, 5904; ISABELA Park, 0 m elevation: KU 300416–30. Platymantis luzo- PROVINCE, Municipality ofPalanan, Barangay Didian, Sitio nensis.—(27) LUZON ISLAND, LAGUNA PROVINCE, Natap Dukan, Northern Sierra Madre National Park, 16u Municipality of Los Ban˜os, Barangay Batong Malake, 579 55.80 N, 122u 249 13.80 E: CAS 204739–41. Platymantis Mt. Makiling: CAS 196364, 196369–70, 200404–08, spelaeus.—(7) NEGROS ISLAND, NEGROS ORIENTAL 210544–45 (Paratypes); CAMARINES SUR PROVINCE, Munic- PROVINCE, Municipality of Basay, Tiyabanan Barrio: CAS ipality of Naga City, Mt. Isarog: FMNH 251643–44; 153477–78, 153482 (Paratypes); NEGROS OCCIDENTAL PROV- TNHC 62004–09, 62012–13, 62020–24; POLILLO IS- INCE, Municipality of Cauayan, Sitio Banso, Barangay LAND, QUEZON PROVINCE, Municipality of Polillo, Bar- Camalandaan, 320 m elevation: KU 300435–38. Platyman- angay Pinaglubayan: KU 305541–42. Platymantis mimu- tis subterrestris.—(3) LUZON ISLAND, MOUNTAIN lus.—(12) LUZON ISLAND, LAGUNA PROVINCE, PROVINCE, Mt Data: CAS 204319–204321. Platymantis Municipality of Los Ban˜os, Barangay Batong Malake, taylori.—(4) LUZON ISLAND, ISABELA PROVINCE, Munic- Mt. Makiling: TNHC 54930–31; PNM 9260–69. Platy- ipality of Palanan, Barangay Didian, Sitio Natapdukan: CAS mantis montana.—(13) LUZON ISLAND, QUEZON 207443–207446 (Paratypes).