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Limnonectes beloncioi, new species urn:lsid:zoobank.org:act: 4CFCEF76-99D6-40DC-9B53- 58DECB1027CF Mindoro Fanged Frog Figures 1, 6, 7 Rana macrodon blythi Boulenger, 1920 (partim). Rana acanthi Taylor, 1923; Taylor and Elbel, 1958. Rana macrodon acanthi (Inger, 1954). Rana macrodon macrocephala (Inger, 1954) (partim: three Mindoro specimens [USNM] provisionally referred by Inger’s ‘‘tentative identification’’ [Inger, 1954: 129]). Rana magna acanthi (Inger, 1958). Limnonectes (Limnonectes) acanthi Dubois, 1987 (partim). Limnonectes cf. acanthi Evans et al., 2003; Setiadi et al., 2010; Diesmos et al., 2015. Holotype.— PNM 9870 (adult male; formerly KU 303343; Field collector No. RMB 4957), Philippines, Mindoro Island, Oriental Mindoro Province, Municipality of Bongabong, Barangay Carmundo, Sitio Paypay-Ama, Paypay-Ama River, 12.7354 ° N, 121.4141 ° E, 100 m above sea level, WGS 84, R. M. Brown, A. C. Diesmos, C. D. Siler, and E. L. B. Rico, 13 March 2005. Paratypes (Paratopotypes).— KU 302084, 302087–88 (adult females), 302085–86, 302089 (adult males), 303343 (juvenile of undetermined sex), 303369–78 (10 subadults of undetermined sex), bearing the same data as the holotype. Other paratypes.— Mindoro Island, Oriental Mindoro Province, Municipality of Bongabong, Barangay Formon: KU 302090–91 (adult females), 302093, 302095, 302097, 302100 (3 adult males, 1 female), 302109–11 (3 adult males), C. D. Siler, 12 March 2005; Municipality of Victoria, Barangay Loyal: KU 302112–18 (2 adult males, 2 adult females, 3 juveniles of undetermined sex), C. D. Siler, 13 March 2005; Barangay Loyal, Sitio Panguisan, Panguisan River: KU 303470–78 (4 adult females, 5 subadults of undetermined sex), R. M. Brown, A. C. Diesmos, and C. D. Siler, 14 March 2005; Municipality of Gloria, Barangay Malamig: KU 302108 (adult female), 303344 (juvenile), J. B. Fernandez and R. M. Brown, 17 March 2006; KU 303346–54 (2 adult males, 2 females, 5 juveniles), R. M. Brown, C. D. Siler, and A. C. Diesmos, 13 March 2005; Sitio Balogbog, Cueba Simbahan: KU 303379–80 (2 subadults of undetermined sex), R. M. Brown, C. D. Siler, and E. L. B. Rico, 12 March 2005; Sitio Pastohan, Tanguisian Falls: KU 303381–402 (22 subadults of undetermined sex), A. C. Diesmos and E. L. B. Rico, 11 March 2005; Occidental Mindoro Province, Municipality of Calintaan, Barangay New Dagupan: KU 303266, 303345 (subadults), R. M. Brown, 8 March 2005; Municipality of Magsaysay, Barangay Nicolas, Sitio Banban: KU 303404–30 (1 adult female, 25 subadults and juveniles /metamorphs of undetermined sex), C. D. Siler and R. M. Brown, 9 March 2005; KU 304131–32 (adult male and subadult of undetermined sex), R. M. Brown; Municipality of Sablayan, Barangay Batong Buhay, Sitio Batulai, Mt. Siburan: KU 303430–52 (5 adult males, 6 adult females, 12 subadults of undetermined sex), E. L. B. Rico, 14 February 2006; KU 305450–51, 306637 (adult female, 2 subadult females), E. L. B. Rico, 19 February 2006; Barangay Malisbong, Sitio Aruyan: KU 335863–83 (11 females, 10 males), S. N. Travers, C. H. Oliveros, and R. M. Brown, 6 July 2013; Barangay Burgos, Sitio Posoy, Posoy River: KU 303453–69 (adult male, adult female, 15 juveniles of undetermined sex), R. M. Brown, 8 March 2005; Municipality of Paluan, Barangay Harrison, Sitio Ulasan, local name ‘‘ Matingaram’ ’: KU 308307, 308309, 308313–18, 308321–23, 308327, 308360, 308362–63, 308367–68, 308370–71, 308385, 308391, 308393, 308422, 308457, 308462, 308464–65, 308469, 308472 (15 adult females, 15 adult males), E. L. B. Rico, 4 January 2007; Municipality of Puerto Galera, Barangay San Isidro, Sitio Minolo, Ponderosa Golf Resort, adult female, J. A. McGuire and V. Yngente, 15 January 1996: TNHC 54920; Municipality of San Teodoro, Barangay Villaflor, Tamaraw Falls, approximately km 15 from Puerto Galera on Calapan-to-Puerto Galera road, 8 subadult males, 1 immature female, 3 adult males, J. A. McGuire and V. Yngente, 17 January 1996: TNHC 54921–29, 55023, 55025, 55029, 55033; same locality, 10 adult males, 11 adult females, R. I. Crombie and V. Yngente, 8 March 1995: USNM 556073–94; Municipality of Baco, Barangay Lantuyan, near Cabinuangang River: 6 adult males, R. I. Crombie and V. Yngente, 2 July 1991: USNM 508558–63; 5 adult males, 3 adult females, 1 immature specimen of undetermined sex, R. I. Crombie and V. Yngente, 7 March 1995: USNM 508564–72; Municipality of Tarogin, ca. 30 km S of Calapan Town, Mt. Halcon SE slope: CAS-SU 22146 (adult female), Q. Alcala and party, 1 April 1963; CAS-SU 22145 (adult female), same data, 31 March 1962; CAS-SU 22147–49 (adult male, 2 adult females), 1 April 1963; CAS-SU 22150 (adult female), S. Magusara and C. Batal, 14 April 1963; CAS-SU 22576 (adult male), Q. Alcala and party, 13 March 1963; CAS-SU 22577, 23508 (adult male and female), 31 March 1963; CAS-SU 23499, 23501, 23525 (adult females), 23505, 23514–15, 23519–20 (adult males), CAS-SU 23485, 23487, 23496–97, 23512–13, 23522 (subadult males), 23489, 23498, 23502 (subadult females), Q. Alcala and party, 10 March 1963; Municipality of Tarogin, Mt. Halcon: CAS-SU 22240 (juvenile), Q. Alcala and party, 14 April 1963; CAS-SU 22288– 22295, 23500, 23510–11, 23517–18, 23521 (juveniles), Q. Alcala and party, 1–20 April 1963; E side of Mt. Halcon, SE slope of Barawanan Peak, 830 m: CAS-SU 22151 (adult female), M. Pinero and party; Semirara Island, Oriental Mindoro Province, Municipality of Caluya, Barangay Tinogboc: KU 302105–07 (2 adult males, 1 adult female), C. D. Siler, 16 November 2000. Referred specimens.— Mindoro Island, Oriental Mindoro Province, Municipality of Baco, Mt. Baco, Alangsa River: USNM 508534–57; Occidental Mindoro Province, Municipality of Paluan, Barangay Harrison, Sitio Ulasan, local name ‘‘ Matingaram’ ’: KU 308308, 308310–12, 308319–20, 308324–26, 308361, 308364–66, 308369, 308372–76, 308386–90, 308392, 308394, 308416–21, 308423, 308430, 308451–52, 308456, 308461, 308463, 308467–68, 308470–87, 308500, 308528, 308538, 308561–69, 308586, 308589, 308590–92; Municipality of Paluan, Barangay 1, Sitio Ipol: KU 308593, 308597, 308599. Diagnosis and comparisons.— Limnonectes beloncioi is a medium-sized fanged frog, assigned to the genus Limnonectes (family Dicroglossidae), on the basis of its prominent, sexually dimorphic odontoid processes characteristic of the genus among other osteological synapomorphies (Inger, 1954, 1966; Emerson and Berrigan, 1993). The new species can be distinguished from all other known congeners based on a combination of its single-pulse/note advertisement call (vs. dual-pulses/note in L. acanthi from Palawan Island faunal region and PAIC landmasses), its phylogenetic position (sister to L. acanthi from Palawan PAIC; Evans et al., 2003; Setiadi et al., 2011), and its geographic distribution on Mindoro and Semirara Islands (vs. Palawan PAIC); it is the only species of Limnonectes known to occur on Mindoro and Semirara Islands and associated small satellite islands and, therefore, has no sympatric congeners. The new species is morphologically similar to its closest relative, L. acanthi; however, it may be distinguished from this allopatric congener by its male advertisement call. The note pulse substructure of L. beloncioi is singular (1 pulse per note vs. 2 pulses per note in L. acanthi), and the new species has a slower note repetition rate (11.5–12.8 notes per second vs. 14.2–15.4 notes per second), and has a higher dominant frequency (2,067.2–2,799.3 Hz vs. 1,335.1–1,679.6 Hz in L. acanthi). With the exception of the morphologically indistinguishable L. acanthi, the new species can be distinguished from all other Philippine species of Limnonectes (L. diuatus, L. ferneri, L. leytensis, L. macrocephalus, L. magnus, L. micrixalus, L. palavanensis, L. parvus, L. visayanus, and L. woodworthi) by a combination of body size, fang (odontoid) length, snout shape, relative lengths of the first and second finger, dorsal skin rugosity, restriction of white-tipped dermal asperities to the sacral region (not aggregated in radial clusters), the presence of irregular, elongate, discontinuous dorsolateral ridges (absence of a continuous dorsolateral fold), complete interdigital webbing of the foot, and the absence of a dark inverted ‘‘V’’-shaped mark on the dorsum. We provide morphological comparisons below, based on our data, in conjunction with or with consideration of the descriptions of Stejneger (1910), Taylor (1923), Inger (1954), Brown and Alcala (1977), and Siler et al. (2009). Limnonectes beloncioi differs from L. diuatus and L. ferneri by its rounded snout in lateral aspect (vs. posteroventrally sloping), its Finger I> Finger II relative finger lengths (vs. approximately equivalent length), and by restriction of white-tipped dorsal asperities to dorsal sacral region, and not distributed in radial clusters (vs. asperities not posteriorly restricted in L. diuatus; and densely distributed across entire of dorsum, and concentrated in radial sacral clusters in L. ferneri); and the presence of irregular dorsolateral ridges (absent in L. diuatus and L. ferneri); from L. leytensis, the new species can be distinguished by its larger adult body size (SVL range 54.2–83.1 vs. 25.8.2–34.0 mm in L. leytensis), rounded snout (vs. snout moderately pointed in lateral aspect in L. leytensis), Finger I> Finger II (equivalent length in L. leytensis), complete webbing (webbing incomplete/reduced in L. leytensis), and the absence of an inverted ‘‘V’’-shaped mark on the anterior dorsum (vs. present in L. leytensis); from a large male specimen of L. macrocephalus (the species endemic to the Luzon PAIC landmasses of Luzon, Polillo, Catanduanes, and Marinduque), the new species can be distinguished by the observation that it attains a considerably smaller maximal adult male body size (SVL range 54.2– 83.1 vs. 78.9–144.6 mm in L. macrocephalus); the new species also possesses a fully exposed tympanum (vs. dorsal and/or posterior edge of tympanum hidden beneath overlapping supratympanic dermal ridge skin in L. macrocephalus) and lacks sexual size dimorphism (Table 2), whereas L. macrocephalus exhibits reverse sexual size dimorphism (males larger); from a large adult male specimen of L. magnus, the new species can similarly be distinguished by its smaller adult body size (SVL range 54.2–83.1 vs. 66.3–164.4 mm in L. magnus), rugose middorsal skin texture (vs. smooth to shagreened in L. magnus), fully exposed tympanum (vs. dorsal and/or posterior edge of tympanum hidden beneath overlapping supratympanic dermal ridge skin in L. magnus), rounded snout (vs. pointed in L. magnus), and by the absence of reverse sexual size dimorphism (present in L. magnus); from L. parvus, L. micrixalus, and L. palavanensis, the new species can be distinguished by its larger adult body size (SVL 54.2–83.1 mm; vs. 24.2–35.5 in L. parvus; 28.1–30.2 in L. micrixalus; 30.0–37.6 in L. palavanensis), rugose (vs. smooth) dorsal skin, Finger I> Finger II (equivalent length), the presence (vs. absence) of white-tipped sacral asperities and the presence of irregular, discontinuous dorsolateral ridges (vs. asperities absent, dorsolateral folds continuous), complete webbing (vs. reduced), and by the absence (vs. presence) of an inverted ‘‘V’’-shaped middorsal marking (vs. present); the new species is additionally distinguished from these species by the absence of sexual size dimorphism (vs. females larger in L. parvus, L. micrixalus, and L. palavanensis); from L. visayanus, L. beloncioi is readily diagnosed by its rounded snout (vs. pointed in L. visayanus), a tendency towards longer adult male fangs (2.6–5.6 vs. 1.8–3.0 mm in L. visayanus), by the presence (vs. absence) of white-tipped sacral asperities; finally, the new species can be distinguished from L. woodworthi by its longer male fangs (2.6–5.6 vs. 1.3– 1.6 mm in L. woodworthi), moderately rugose middorsal skin (vs. smooth in L. woodworthi), rounded snout (vs. snout moderately pointed in L. woodworthi), the presence (vs. absence) of white-tipped sacral asperities, and by the absence (vs. presence) of continuous dorsolateral folds. Description of holotype.— A mature male, specimen in excellent condition; small portion of liver preserved separately for genetic material; habitus robust; head broader than body, its length 98.3% of its width, 39.0% of SVL; snout tip rounded in dorsal and lateral aspect (Fig. 6); supralabial region markedly swollen, increasingly protuberant towards angle of jaw; interorbital region and dorsal rostrum nearly flat; eye diameter 62.0% snout length, 97.5% eye–nares distance, 1.4 ⅹ eye–tympanum distance; pupil horizontally sub-elliptical with discontinuous posterior margin; canthus rostralis distinct, slightly medially bowed in dorsal aspect; loreal region concave; nostrils oriented dorsolaterally, narial openings visible in dorsal view; internarial region slightly convex; tympanum exposed, annulus slightly distinct, diameter 55.7% of eye diameter; supratympanic fold thick, strongly protuberant, moderately rugose, extending from posterior corner of eye, extending horizontally over (concealing) dorsoposterior corner and posterior margin of tympanic annulus, turning ventrally at nearly right angle, to end in supra-axillary region, where it is discontinuous with postrictal tubercular swelling at angle of jaw. Tongue elongate, tapered anteriorly, with narrow anterior attachment and laterally expanded, free, bilobed posterior margin at rest (anterior edge when tongue projected); choanae situated at anterolateral edge of palate, subcircular, their anterolateral edge partially concealed by palatal shelf of maxilla in ventral view; choanae widely separated by distance five or six times greater than diameter of single choana, each located just anterolaterally to (in contact with) lateral tip of dentigerous process of vomer; dentigerous process of vomer distinct, with four or five conical teeth on each side; dentigerous process angled anterolaterally (rostrally), approximately at 45 ° inclination, with closest (posterior) points separated by distance approximately equal to one choana, their most distant (anterior) ends separated by distance equal to three choanae; enlarged odontoid ‘‘fangs’’ large, recurved, unsheathed by oral mucosa for> distal half their length, situated on either side of mandibular symphysis/medial bulge, their tips sharply pointed, total length 4.6 mm (perpendicular distance from ventral edge of mandible), inclined dorsoposteriorly, tips 2.9 mm perpendicular from dorsal mandible surface; maxillary fang ‘‘sockets’’ anteromedial to choanae, large, round, similar in size to one choana; vocal apertures large, elongate, surrounded by extensive mucosal invaginations, situated at posteroventral margin of buccal floor, just medial to angle of jaw. Hand length 46.8% foot length; foot 93.4% tibia length; tibia length 58.3% SVL; fingers laterally, irregularly ovoid in cross section, due to presence of slight lateral dermal flange, extending from base of each digit, on either side, to proximal margins of terminal finger discs; terminal discs not expanded beyond widths of penultimate phalanges (Fig. 6), their relative descending lengths: III> I> II ¼ IV; subarticular tubercles prominent, their ventral surfaces convex and velvety in texture; one subarticular tubercle below Fingers I and II, two tubercles under Fingers III and IV; terminal discs and subarticular tubercles with gray, velvety, thickened surfaces; distal margins of tubercle more distinct than their proximal margins, and supernumerary tubercles absent, but articular surfaces of fingers between subarticular tubercles of digits, and at base of all digits covered medially with thickened tubercular surface; palmar surface with large, elongate, thenar tubercle (ventromedial surface of Finger I), enlarged, flattened, squarish medial ‘‘inner’’ (base of Finger III) metacarpal tubercle, and small, ovoid, convex outer (base of Finger IV) metacarpal tubercle; surface of these palmar structures, intervening, and surrounding surfaces all covered with similar, thickened, velvety (matte) tubercular dermis layer; nuptial excrescences or pads, asperities, and webbing absent; forearm musculature not hypertrophied. Tarsus folds and flaps absent; terminal discs of toes slightly expanded, with distinct circummarginal grooves; plantar surfaces of foot with well-developed, prominently protruding (ventrally), rounded subarticular tubercles (Fig. 6); plantar surfaces of foot smooth, with velvety-textured subarticular tubercles; relative lengths of toes: I Skin of dorsal surfaces of trunk and head smooth to slight shagreened texture, bearing low but clearly evident fleshy dermal tubercle clusters in supra- and post-tympanic regions, and tuberculate dorsolateral ridges, immediately following supratympanic region, and continuing posteriorly to the scapular region approximately to the points of forearm insertion (Fig. 6); similar dorsolateral tubercular ridges extend from this point, along more lateral (flanks) and slightly medial (dorsal) surfaces, and extend back to the sacral region; on posterior half of trunk and sacral region, dermal tubercles present mid-dorsally, consisting of single, raised tubercles or short, raised tubercular ridges; in sacral region, some tubercles capped with round, weakly keratinized dermal asperities; not arranged in clusters, or rows, but lightly dispersed in sacral region and upper one-third of dorsal surface of thigh and supra-cloacal region; ventral surfaces of head smooth; lateral and ventral surfaces of limbs smooth; remaining dorsal surfaces of limbs smooth to lightly shagreened, with occasional low tubercles; tarsus smooth on dorsolateral surface; cloacal region rugose (wrinkled), with smooth laterally and ventrally surrounding skin. Coloration of holotype in preservative.— Dominant dorsal color on head, body, and forelimbs uniform Dark Grayish-Brown (Köhler, 2012; color 284) Dark Grayish-Olive (275) with irregular, diffuse, Dusky Brown markings (285) concentrated on occiput and sacral regions; lateral head surfaces Drab (19) with diffuse Warm Sepia (40) m

Herr, Mark W., Goyes Vallejos, Johana, Meneses, Camila G., Abraham, Robin K., Otterholt, Rayanna, Siler, Cameron D., Rico, Edmund Leo B., Brown, Rafe M. (2021): A New, Morphologically Cryptic Species of Fanged Frog, Genus Limnonectes (Amphibia: Anura: Dicroglossidae), from Mindoro Island, Central Philippines. Ichthyology & Herpetology 109 (1): 188-210, DOI: 10.1643/h2020095, URL: http://dx.doi.org/10.1643/h2020095

Chan, Kin Onn, Abraham, Robin K., Grismer, L. Lee, Brown, Rafe M. (2020): A systematic review of the Pulchrana picturata complex, with the description of a new species from Peninsular Malaysia, Sumatra, and southern Thailand. Raffles Bulletin of Zoology 68: 880-890, DOI: 10.26107/RBZ-2020-0096

Cornufer (Batrachylodes) exedrus sp. nov. Figs. 2a–c, 4, 5 Holotype. South Australia Museum (SAMA) R64760 (SJR Field No. 10784), adult male, collected by S. J. Richards at Tompoi Camp, 1590 m above sea level (05°20.623'S, 151°18.873'E; WGS-84), Nakanai Mountains, East New Britain Province, Papua New Guinea, 19 April 2009. Paratypes. SAMA R64762 and R64764 (SJR Field Nos. 10815 and 10820 respectively), two adult males, and SAMA R64763 (SJR 10819), adult female, all collected at the type locality, 21 April 2009; SAMA R64761 and R71008 (SJR 10798 and 10821 respectively), two adult males, same locality, 20 and 21 April 2009, respectively; SAMA R64765 (SJR 10843), juvenile, and SAMA R71007 (SJR 10846), adult male, same locality, 23 April 2009. Etymology. The specific epithet is derived from the masculine formation of the Greek adjective exedros, meaning away from home, strange, or extraordinary. The surprising discovery of Cornufer (Batrachylodes) exedrus from a high elevation site in New Britian is a taxonomic and biogeographic enigma, given that all other species in the subgenus are restricted to the Solomon Archipelago. Diagnosis. The new species is a member of the genus Cornufer based on molecular phylogenetic evidence. There are currently no morphological synapomorphies known for the genus Cornufer, however, based on the updated taxonomic arrangement of this clade (Brown et al. 2015) into several morpholgically diagnosible subgenera the new species can be assigned to the subgenus Batrachylodes, as defined by Brown et al. (2015), on the basis of its small, triangular shaped body, pointed snout, absence of interdigital webbing, absence of vomerine teeth, presence of a darkened loreal stripe extending diagonally across the flanks, and molecular phylogenetic placement (Brown et al. 2015:fig. 2). Cornufer exedrus is distinguished from congeners by a combination of (1) adult body size small (SVL 17.6–18.8 mm for six adult males, mean = 18.3 mm; SVL 18.3 mm for one adult female), (2) body shape triangular with a small, pointed head, (3) terminal disks of the fingers and toes narrowly expanded (among adults: Fin3DW 0.3–0.6 mm, mean = 0.5 mm; Toe4DW 0.4–0.7 mm, mean = 0.6 mm) (Figs. 2, 4, 5), (4) vomerine teeth absent, (5) digits lacking all vestiges of lateral flange and interdigital webbing, (6) dorsal skin smooth to finely granular, lacking prominent dermal tubercles or ridges, (7) subarticular tubercles of fingers, toes, hands and feet uniformly low and flattened, (8) dark band extending along the canthus rostralis and onto the flank with a smooth dorsal margin demarcating lighter dorsal and darker lateral colors, and (9) unique advertisement call consisting of a series of brief, high-frequency atonal notes. Comparisons with other species. Among members of the subgenus Batrachylodes, Cornufer exedrus can be easily distinguished from C. elegans, C. gigas, C. montanus, C. vertebralis, and C. wolfi by the narrowly expanded (versus widely expanded) terminal finger disks except for the first finger (Fin3DW/SVL = 0.02–0.03, Fin3DW/ Fin3L = 0.1–0.2, and Fin3DW/TD = 0.2–0.3 in C. exedrus versus Fin3DW/SVL> 0.03, Fin3DW/Fin3L> 0.2, and Fin3DW/TD> 0.3 in the aforementioned species) and smaller adult body size (SVL 17.6–18.8 mm in C. exedrus versus SVL> 20.0 mm in the aforementioned species) (Table 2). Of the Batrachylodes that lack prominently dilated finger disks, Cornufer exedrus differs from C. mediodiscus in having a smaller adult body size (SVL 17.6– 18.8 mm in C. exedrus versus 20.2–26.0 mm in C. mediodiscus), slightly smaller third finger disk width (reflected in the Fin3DW/SVL, Fin3DW/Fin3L, and Fin3DW/TD ratios, Table 2), and lacking raised dorsal skin ridges (Brown & Parker 1970). Cornufer exedrus differs from C. minutus by its slightly larger body size (SVL 17.6–18.8 mm in C. exedrus versus 15.2–17.3 mm in C. minutus) and by differences in the proportions related to hind limb length (reflected in the HW/TBL, TBL/SVL, and PL/SVL ratios, Table 2). Cornufer minutus also differs in color pattern, with the dark band along the canthus rostralis more poorly defined, often broken or with an irregular dorsal margin, than in Cornufer exedrus. The new species differs from C. trossulus by its slightly smaller body size (SVL 17.6–18.8 mm in C. exedrus versus 18.9–24.3 mm in C. trossulus) and by differences in the proportions related to hind limb length (reflected in the HW/TBL, TBL/SVL, and PL/SVL ratios, Table 2). On New Britain Island, C. exedrus can be differentiated from most congeners by its smaller adult body size (Cornufer species on New Britain (C. exedrus can be further distinguished from C. caesiops (and the morphologically similar C. browni, on nearby New Ireland Island; Allison & Kraus 2001; Kraus & Allison 2009) in having narrowly expanded finger disks (versus widely expanded in C. caesiops and C. browni); from C. akarithymus, C. bufonulus, and C. sulcatus in having a smooth dorsal surface (versus tuberculate or with dorsal ridges in C. akarithymus, C. bufonulus, and C. sulcatus); and from C. boulengeri (males and juveniles previously assigned to C. rhipiphalcus; Kraus 2008), by the lack of a fanlike array of dorsal ridges in the scapular region and a less prominent dark face mask, which does not extend onto the flanks in C. boulengeri. Description of holotype. Adult male, in good condition, preserved with mouth open, incision in left thigh where SAMA tag was threaded; habitus moderately robust; head not distinct, slightly narrower in dorsal aspect than body, length 41% of SVL; head length 120% of head width; snout short, tip acuminate in lateral aspect, rounded dorsal aspect, protruding prominently beyond lower jaw, unpigmented at tip; lower jaw notched at tip; eyes protrude only slightly beyond silhouette of head in both dorsal and lateral aspects; labial region rounded and smooth, lips not swollen, not extending past eyes when viewed in dorsal aspect; interorbital region slightly convex; eye diameter 90% of interorbital distance; pupil horizontally ovoid; canthus rostralis medially bowed; loreal region slightly concave; eye diameter 55% of snout length; nostrils slightly laterally protuberant; eye-narial distance 1.25 times the distance from nostril to snout tip; internarial region flat; tympanum distinct, its diameter 89% of eye diameter; slight supratympanic fold present, extending from posterior edge of eye, over dorsal edge of tympanum, and terminating at supra-axillary (post-rictal) region; tongue ovoid, with a slightly narrower anterior attachment; choanae round, minute, at anterolateral edge of palate, separated by a distance five to seven times greater than their diameter, obscured in ventral aspect by palatal shelf; vomerine teeth absent; openings to vocal sac minute slits, at level of the angle of jaw. Dorsal skin of body, head, and limbs generally smooth, lacking dermal folds, tubercles, or ridges; ventral surfaces of trunk, head, throat, and limbs smooth, with the exception of fanlike array of 12–14 small white tubercles on anterior margin of chin. Cornufer Batraychylodes......continued on the next page 9 5: 6;!#") C. montanus C. gigas & Manus length 50% of foot length; digits of manus narrow (Figs. 4–5), ovoid in cross-section; dermal flanges absent on lateral edges of digits; terminal finger disks barely expanded ( Hindlimbs short; tibia length 41% of snout-vent length, pes length 99% of tibia length; tarsus smooth, lacking folds, flaps, or tubercles; terminal disks of toes slightly expanded and ovoid ( Measurements of holotype. SVL 18.1; ED 1.8; TD 1.6; HL 7.4; SNL 3.3; IOD 2.0; HW 6.2; FL 8.1; TBL 7.5; TSL 4.5; PL 7.4; ML 3.7; FA 3.7; Toe4L 4.1; Fin1L 1.1; Fin3L 2.2; Fin1DW 0.3; Fin3DW 0.3; PpFin3 0.2; Toe4DW 0.4; PpToe4 0.3. Color of holotype in preservative. Dorsal surfaces of body and head exhibit nearly homogeneous light brown coloration; under magnification, background color of dorsal surfaces is pale cream and unpigmented with dense and nearly uniform speckling of brown chromatophores; less brown pigmentation present in interorbital region, extending posteriorly in a U-shaped pattern to a midpoint between the scapulae, giving slightly paler appearance to region along top of head than rest of body; two small, dark brown interorbital spots between anterior margin of eyes; tip of snout unpigmented; dorsal surface of limbs similar in background coloration to rest of body, but with darker brown transverse bars (three on forearm, three on femur, three on tibia, one on tarsus, two on metatarsus) with irregular margins and poorly defined; dorsal surfaces of hands, feet, and digits similarly colored as rest of the body and limbs with irregular dark spots and bars; the lateral surfaces of the head and body have a distinct dark brown band (darker than dorsal coloration) covering entire loreal and supralabial region, extending posteriorly along flanks diagonally (anteriodorsally to posteroventrally) towards inguinal region; this lateral band is darkest and near uniformly shaded along head (although tympanum is slightly lighter) becoming lighter and more diffusely speckled posterior to the humerus, into light tan irregular blotching in the inguinal region; the dorsal margin of this lateral band is nearly smooth and darkest brown in coloration (with respect to the rest of band), forming a clear dark line demarcating lighter dorsal and darker lateral colors extending from snout tip (although absent from unpigmented snout tip) along the canthus rostralis (above nostril) and upper eyelid, and posterior onto flanks slightly above tympanum; dark dorsolateral line fades in the inguinal region becoming prominent again on hindlimbs (although with more irregular margin) particularly on anterior edge of thigh and knee, fading into diffuse irregular tan blotching on tibia; underside of throat dark brown with lighter unpigmented mottling throughout; brown ventral coloration of throat fades posterior to the pectoral region becoming diffuse irregular tan blotching on pale cream midventer; ventral surfaces of forearms tan, darker brown towards outer edge of forearms; inner forearm with unpigmented band extending to underside of humerous; underside of hindlimbs similar in coloration to midventer, pale cream with brown irregular mottling; when viewed in posterior aspect cloacal region nearly uniform dark brown with smooth, dark dorsal margin above cloaca extending along the posterior of thighs clearly demarcating it from lighter dorsum; when in a seated position, this dark cloacal-femoral band grades into similarly colored dark band along outer edge of tarsus; small dark brown blotch just above cloaca; ventral surfaces of hands and feet dark brown with pale subarticular banding. Variation. Compared to the holotype (SAMA R64760), dorsal coloration of the head and body of the paratypes ranges from a darker brown (SAMA R64763, SAMA R64764, SAMA R71008, SJR 10843) to a lighter cream (SAMA R64761, SAMA R64762, SAMA R71007) background coloration, and whereas the holotype has a nearly uniform dorsal coloration, all of the paratypes exhibit some degree of dorsal markings. SAMA R71008, SAMA R64763, and SJR 10843 possess a faint, thin and lighter vertebral stripe. Four of the paratypes (SAMA R71007, SAMA R71008, SAMA R64762, SAMA R64761) possess a pair of discrete darker brown to black dorsal spots that are posteroventrally located just behind the projection of the sacrum. Several paratypes also have irregular darker paravertebral spots between the scapulae, which are similar in size to the posterior dorsal spots in some (SAMA R64761, SAMA R64762) or a faint speckling in others (SJR 10843, SAMA R64763). SAMA R64761 exhibits more extensive irregular dark flecking and blotching along the dorsum. Three paratypes (SAMA R64764, SAMA R64761, SAMA R71007) possess a faint, thin and darker interorbital bar. Several paratypes also exhibit faint darker banding extending from behind eyes and/or the sacral region intersecting at the midscapular region, leaving a faint x or v-shaped pattern along the dorsum. All of the paratypes possess the transverse bars on the limbs as described in the holotype. There is variation in breadth and extent of the dark lateral band that extends from the loreal region onto the flanks. In some individuals, the lateral band is darker and more filled in throughout the snout and flank region, giving appearance of dark face mask which remains solid (not fading) along the flanks. However, in others the lateral band is limited to more of a dark stripe from the tip of the snout, along the canthus rostralis, and posterior towards the inguinal region, which quickly fades ventrally into a light-brown coloration similar to the color of the dorsum. In two individuals (SAMA R64761, SAMA R71007), the lateral band is almost entirely limited to the snout along the canthus rostralis and quickly broken and fading posteriorly behind the eye, and in these individuals there is almost no darker shading below the band in the loreal and supralabial region of the snout or along the flanks. Ventral coloration of several of the paratypes is similar to that of the holotype, with a darker brown chin and throat region fading into a lighter cream midventral region and the underside of the hindlimbs with diffuse brown speckling. However, in one individual (SAMA R64762) the entire vertrum is similar in coloration to the brown throat, and in three individuals (SAMA R64761, SAMA R71007, SJR 10843) the brown pigmentation has faded, leaving the entire ventrum pale cream in coloration. In most of the paratypes the skin texture is smooth, similar to that of the holotype, however several specimens exhibit finely granular skin along the dorsum, and one indiviudal (SAMA R64761) possesses small raised dorsal ridges and minute tubercles associated with the darker brown spots and flecks along the dorsal surface of the body. The small unpigmented tubercles on the chin and throat of the holotype are also present on most of the paratypes (although varying in number and extend along the throat); however, they are absent on the juvenile (SJR 10843) and female (SAMA R64763) specimens, suggesting that it may be a sexually dimorphic character. Additionally, the unpigmented projection of the snout on the holotype is prominent on most of the paratypes except the aforementioned juvenile and female paratypes, suggesting it may also be a sexually dimorphic character. Mensural variation is presented in Table 2. All mensural ratios for the single female paratype are within the ranges found in males, possibly indicating little sexual dimorphism in morphometric characters. Color in life. Based on color images of the holotype (SAMA R64760, Figs. 2–3) and two paratypes (SAMA R64763 and SAMA R64764, Fig. 2) in life taken during the daytime, prior to preservation: Dorsal ground coloration of body and limbs of the holotype (SAMA R64760) a pale pinkish-salmon that is overlain with small darker pinkish-orange spots or streaking (streaks most prominent on the tibia) with some small irregular whitish spotting throughout; thicker (but faint) darker brown transverse bars extending across the hindlimbs and three thin (broken) darker brown transverse bars on the forearm; dorsal coloration of the head orangish, giving the appearance of a head cap (from just behind the unpigmented snout projection extending in a U-shaped pattern posterior to the eyes to the mid-scapular region); three small whitish and two small dark brown interorbital spots between the anterior margins of the eyes; margin of the upper eyelid with a thin light whitish band speckled with diffuse brown flecking. Dorsal coloration of the two paratypes different than that of the holotype. One paratype (SAMA R64763) has a more solid and uniformly darker brown coloration across the head, body, and limbs flecked with minute irregular dark brown, whitish, and reddish spots; limbs also with some suffused lighter orangishbrown highlights; a thin and slightly broken vertebral stripe is present, and a cluster of small dark-brown paravertebral spots in the midscapular region; faint darker brown transverse bars present on the limbs. The other paratype (SAMA R64764) possesses lighter tan dorsal coloration on the head and body, fading to darker brownishgray hindlimbs (forearms of similar color as hindlimbs), leaving the transverse bars on the hindlimbs poorly diagnosable; faint minute irregular dark-brown, whitish, and reddish spots or blotching on the dorsal surfaces of the body and limbs; light yellow-gold pigment faintly dusting the forelimbs and hands; two darker brown dorsal bands also apparent, one interorbital band between the midpoints of the orbits and upper eyelids, and one extending as a forward-facing V-shape connecting at the midscapular region and extending posteroventrally to the mid-dorsum. The shading of the dark-brown lateral band (from the snout tip to the flanks) also varies slightly. In all three specimens the dorsal margin of the lateral band is the darkest brown, particularly in the loreal region along the canthus rostralis; in SA

Platymantis quezoni, sp. nov. ��� Platymantis sp. 27 ��� Brown et al. 2015 Figs. 2���4 Holotype. PNM 9817 (RMB 4067; formerly KU 339542), adult male, collected on limestone outcrop in original forest (19:00 hr) on 27 November 2001 in Barangay Malinao Ilaya, Municipality of Atimonan, Quezon Province, Luzon Island, Philippines (N 13.989 ��, E 121.818 ��; 275 m elevation; datum: WGS 85), by MLLD and G. V. A. Gee. Paratypes (paratopotypes). Sixteen males: TNHC 61989 ��� 94, bearing the same data as the holotype; PNM 9910 ��� 12, collected on 19 October 2013 by LADL, AL, MLLD, and Jason Fernandez; PNM 9913���9916, collected on 15 December 2014 by LADL and AL; KU 328735 and 328736, bearing the same data as the holotype; KU 339543, collected 9 January 2011; three females (PNM 9907 ���09, collected on 19 October 2013 by LADL, AL, MLLD and Jason Fernandez; one juvenile (KU 339544, bearing the same data as the holotype), all collected at the type locality. Diagnosis. Platymantis quezoni, sp. nov., can be distinguished from other Philippine congeners by the following suite of characters: body size 22.1���33.9 mm SVL for males, 33.8���39.7 mm for females; terminal discs of fingers and toes slightly expanded (Fig. 2 A,B); Finger I longer than Finger II; fingers and toes lacking dermal flanges along sides of digits, subarticular tubercles large and highly protuberant (Fig. 2 A,B); skin smooth, but with scattered and raised dermal tubercles (Figs. 3, 4); dorsolateral folds, mid-dorsal ridges, and other cephalic protuberances absent; color pattern medium to dark brown with black spots (Fig. 3); advertisement call a series of invariant low-frequency stridulated one-syllable barks: ���Iyohk, iyohk, iyohk������ (Fig. 6) and its restriction to terrestrial microhabitats in forest covering limestone karst formations (Fig. 5). Diagnostic character state differences are summarized in Table 1 and we provide detailed comparisons, emphasizing diagnostic differences between the new species and its close relatives below. Comparisons. We assign the new species to the subgenus Lupacolus Brown et al. 2015 on the basis of its placement (as ��� Platymantis sp. 27 ���) in Clade R of the genus Platymantis (Brown et al. 2015: Fig. 2). The subgenus Lupacolus is largely equivalent of the former P. dorsalis Group of Brown et al. (1997 a) and Alcala & Brown (1999), and contains the named taxa P. cagayensis, P. dorsalis, P. indeprensus, P. mimulus, P. naomiae, P. paengi, P. pseudodorsalis, P. spelaeus, and P. t a y l o r i but does not include P. biak, P. levigatus, P. pygmaeus (placed in subgenus Lahatnanguri; Clade O of Brown et al. 2015: Fig. 2) or P. corrugatus (subgenus Platymantis, clade name Tagomukhus; Clade N of Brown et al. 2015: Fig. 2). Members of the genus Platymantis, subgenus Lupacolus, all possess the following combination of characters: body size moderate (~ 25���45 mm SVL in males); digital discs non-expanded to minimally expanded; first finger longer than second; fingers and toes lacking dermal flanges along sides of digits; subarticular tubercles large and highly protuberant (Brown et al. 1997 a, c; Alcala & Brown 1999; Fig. 3). In overall phenotypic similarity Platymantis quezoni most resembles the related species P. paengi, P. spelaeus, (all three belong to the subgenus Lupacolus), the unrelated P. biak, P. insulatus (both of the subgenus Lahatnanguri), and P. bayani (subgenus Tahananpuno). All six taxa share the general mottled limestonecamouflaged color pattern and all exclusively inhabit limestone karst microhabitats. Platymantis quezoni shares with the other members of the subgenus Lupacolus (P. paengi, P. spelaeus) minimally expanded terminal digital discs, whereas these two species of subgenus Lahatnanguri (P. biak, P. insulatus), and all members of the subgenus Tahananpuno (including P. bayani) have widely expanded terminal discs of the fingers and toes (Brown & Alcala, 1970; Brown et al. 1997 a, c; Siler et al. 2009, 2010). Additionally, P. bayani, P. insulatus, and P. spel aeus further differ by their much larger body sizes, which do not overlap the size range of P. quezoni (Table 1). Finally, the advertisement call of the new species (a rapid series of loud, harsh, single-syllable barks) differs from that of P. biak (a much slower, low amplitude series of tonal notes), P. insulatus (a long clicking train of brief atonal pulses), P. spelaeus (a four-syllable, multinote complex call) and P. paengi (tonal, frequency sweep, amplitude modulated call). Platymantis quezoni is distinguished from the remaining members of the subgenus Tahananpuno (P. diesmosi, P. luzonensis, P. negrosensis, P. guentheri, and P. rabori) by the presence of minimally expanded (vs. widely expanded) terminal digital discs of the fingers and toes. The terrestrial limestone karst microhabitat of P. quezoni further distinguishes the new species from all of these arboreal tree frog taxa except P. d i e s m o s i (a terrestrial species calling exclusively from cliff ledges on Mt. Malino; Brown & Gonzales 2007) and its advertisement call likewise distinguishes P. quezoni from the tree frogs with amplitude modulated calls (P. luzonensis, P. negrosensis, and P. rabori) and P. guentheri, an arboreal species with a tonal, frequency sweep call. The new species is distinguished from all species of Philippine cloud frogs (formerly the P. ha z e l a e Group species) of the subgenus Tirahanulap (P. hazelae, P. isarog, P. lawtoni, P. montanus, P. panayensis, P. polillensis, P. sierramadrensis, and P. subterrestris) by its larger body size (vs. much smaller in these species, typically ~ 20���28 mm in males), by unadorned digits (vs. digits with wide dermal fringes), by strongly protuberant (vs. ventrally flattened) subarticular tubercles, by possessing a longer first finger than the second (vs. second longest) and its terrestrial limestone karst microhabitat (vs. shrub-layer vegetation of montane forests). Species of the subgenus Tirahanulap all possess tonal, constant frequency calls with complex harmonic structure, contrasting sharply with the loud, harsh barking calls of P. quezoni. From remaining members of the subgenus Lupacolus (previously referred to as the Platymantis dorsalis Group) (P. cagayensis, P. dorsalis, P. indeprensus, P. mimulus, P. naomiae, P. pseudodorsalis, and P. tayl ori), P. quezoni is distinguished by from all species except P. levigatus by the absence of dorsal folds and/or ridges, and limitation of dermal tubercles to its eyelids (vs. presence dermal folds/ridges and widely distributed tubercles over most dorsal body surfaces), and from all Lupacolus except P. paengi and P. s p e l a e us, by its terrestrial limestone forest microhabitat (vs. much more generalist inhabitant of leaf litter of forest floor and/or herb-layer vegetation in P. cagayensis, P. dorsalis, P. indeprensus, P. mimulus, P. naomiae, P. pseudodorsalis, and P. t a yl or i). Other than P. dorsalis and P. pseudodorsalis (which possess frequency sweep calls), these species possess complex, multisyllable call structure, which contrasts markedly with the simpler, single-syllable barking calls of P. quezoni. The new species differs from Platymantis corrugatus (subgenus Platymantis, clade name Tagomukhus) by its smooth to shagreened skin texture (vs. dorsal skin with elongate dermal folds), by its lateral head coloration similar to other dorsolateral surfaces (vs. presence of a dark ���mask��� coloration on the lateral side of the head of P. corrugatus) and by its exclusive limestone karst microhabitat (vs. a variety of terrestrial forest types). The advertisement call of P. quezoni is surprisingly quite similar to that of P. corrugatus (a loud series of single ���quacks���) except that P. corrugatus is generally crepuscular whereas P. qu e z o ni calls beyond sunset and well into night-time hours if precipitation is sufficient. The final set of comparisons for the recognition of P. quezoni is to the group of species comprising a recently discovered clade for which the subgenus Lahatnanguri was established (Brown et al. 2015). The new species differs from tree frogs of this clade by its terrestrial limestone karst microhabitat and possession of minimallyexpanded terminal discs of the fingers and toes (vs. widely expanded discs in the arboreal species P. banahao, P. cornutus), from the forest floor leaf litter-inhabiting, miniature species P. pygmaeus (SVL 13���16 mm in males) by its much larger body size (22���34 mm) and limestone karst microhabitat, and from the semi-aquatic species P. levigatus by its light brown, mottled dorsal coloration (vs. dark, more uniformly colored dorsum), by its drier, limestone karst microhabitats (vs. moist, riparian environments). Finally, P. quezoni differs from the limestone karst frogs of the subgenus Lahatnanguri by its minimally-expanded terminal discs of the fingers and toes (vs. markedly expanded in P. biak, and widely expanded in P. insulatus), the presence of distinct transverse banding patterns of the thigh and tibia (vs. absent or vague in P. insulatus), and its simple, single-syllable barking calls advertisement call (vs. atonal clicking pulses in P. insulatus and the slow series of low, tonal notes in P. biak.) Description of holotype. A mature male (Fig. 3); habitus moderate; head as wide as body, head length 43.9 % snout���vent length; snout protruding moderately beyond lower jaw, snout tip moderately pointed in dorsal and lateral aspects; snout 39.1 % head length; eyes moderate and anterolaterally protruding, beyond width of head in dorsal aspect; eye diameter 15.1 % snout���vent length, 34.4 % head length; lips flared, labial region slightly swollen (Fig. 4 A); interorbital region flat; eye diameter 87.9 % snout length, eye���naris distance 74.5 % eye diameter; pupil horizontally elliptical; canthus rostralis strait, only sightly medially bowed; loreal region moderately concave, sloping ventrolaterally to labia; nostrils oriented laterally; internarial region slightly concave; tympanic annulus distinct, 60.1 % eye diameter; supratympanic fold moderately protuberant, swollen and thick, extending from posterior corner of eye across entire dorsal margin of tympanum, along postero-dorsal edge of annulus, and to supra-axillary region. Tongue elongate, wider posteriorly, with shallow posterior notch, a single small anterior lingual papilla, and narrow anterior attachment; choanae situated at anterolateral edge of palate, irregularly ovoid, separated by distance six greater than diameter of single choana, partially concealed in ventral aspect by maxillary shelf; dentigerous process of vomer distinct, with four teeth visible on left process, six on right; dentigerous process angled slightly anterolaterally, approximately at 20���30 �� incline with closest (posterior) points separated by distance equal to 2 X diameter of single choana, their most distant (anterior) ends separated adjacent choana by a distance equivalent to one choana. Hands and feet large; manus length 52.6 % pes length; limbs well developed; tibia length 63.4 % snout���vent length, 111.9 % pes length; fingers slender, long; terminal discs moderately expanded (Fig. 2 A) with circummarginal grooves evident around distal edge of terminal discs; dorsal surfaces of terminal phalanges with small, cutaneous supra-articular flaps between ultimate and penultimate phalanges; relative lengths of fingers II Measurements of holotype (in mm). Snout���vent length 33.7; head length 14.8; head width 13.9; snout length 5.8; upper arm length 5.9; forearm length 7.5; femur length 18.1; tibia length 21.7; tarsus length 10.9; pes length 19.4; manus length 10.2; Finger I length 5.9; Finger III length 7.7; Toe IV length 12.6; eye���narial distance 3.8; interorbital distance 3.2; internarial distance 3.0; eye diameter 5.1; horizontal tympanic annulus diameter 3.1; Finger I disc width 1.0; Finger III disc width 1.2; Toe IV disc width 1.3; Finger III penultimate phalange width 0.8; Toe IV penultimate phalange width 0.7. Coloration of holotype in preservative. Dorsal surfaces of head, body, and limbs dark brown with black blotches; anterior portions of head darker greenish brown; limbs banded with black bars; interorbital bar present but faint: a thin, irregular, black line; lateral surfaces of head darker than dorsum, nearly greenish brown with black markings; black bar extends across dorsal half of canthus rostralis to anterior edge of eye, around margin of the palpebrum, and across suptratympanic fold; tympanum medium brownish-green with black covering dorsal-most half; lips with alternating dark green and black vertical labial bars, bordered anteriorly by black lower edge of canthal and subocular regions; three bold black bars radiate out of orbit dorsally, and extend ventrally to labial region to form the most conspicuous labial bars; transverse limb bars of irregular widths, with an overall darker coloration (compared to dorsum), due to a greater proportion of surface covered with black markings; upper arm almost devoid of black blotches, forearms each with two transverse black bars, thighs with five, tibia with three, tarsus with two; anterior flank coloration similar to dorsum on left, but predominantly black on right due to aggregation of black blotches, both sides progressively fading to less strongly patterned ventral color: dark brown blotches on throat, sternal region tan, inguinal region tan with steel bluish markings; ventral surfaces of limbs solid dark reddish brown; ventral surfaces of hands and feed dark reddish brown with light cream to tan subarticular tubercles; dorsal surfaces of hands are light brown, with alternating bands black bars on Fingers II���IV, and light tan on Finger I; dorsal surfaces of feet with solid redish brown above Toes I���III, boldly alternating light brown and black on Fingers IV and V. Coloration in life. Dorsal surfaces of head, body, and limbs variably mottled shades of dark brown and greenish-gray, a few scattered bright gold spots, and with bold black markings scattered throughout but congregating irregularly into the following characteristics: distinct black suprascapular bar, interorbital bar, transverse limb bars (on thigh, tibia, shank, and upper arm and forearm), a dark bar from the nares, along the canthus, aligned with the enlarged pupil at night (Fig. 3), and posteriorly across the dorsal-most half of the tympanum; upper lips with distinct black labial bars; pupil bordered by bright yellow ciliary ring, iris silver; flanks with enlarged black blotches and interrupted by reduced dark brown of dorsal surfaces, grading ventrally across flank to cream ground color with black blotches on anterior portion of flanks; dorsal surfaces of digits mostly black, with small brown spots. Ventrum very dark gray with black blotches on throat and sternum; liver nearly bluish gray through translucent ventral skin of chest and groin; ventral surfaces of legs nearly solid black, fading to dark purplish gray on ventral tibia and shank; ventral surfaces of hands and feet dark purplish gray with lighter gray subarticular tubercles and distal digital discs. Variation. Overall color pattern consistently mottled across the type series, typical of limestone karst inhabiting Platymantis (Brown & Alcala, 1970, 1982a; Siler et al., 2009, 2010), with dorsal ground color light tannish yellow or mustard (e.g., KU 328735; Fig. 3) to dark greenish-gray with black blotches (the holotype: PNM 9817; Fig. 3). The degree and concentration of dorsal black markings ranges, from specimens nearly covered dorsally in black markings (KU 339543, TNHC 61993), to individuals predominantly light gray to tan, with few black markings (KU 61989, 61991, 61992; Fig. 3 B). The suprascapular bar ranges from a distinct w-shaped pattern (the holotype: PNM 9817; TNHC 61990; KU 328736) to a few irregular, disconnected blotches (TNHC 61991) to an inverted v-shaped formation (KU 339543). Ventral surfaces less variable, predominantly as described for the holotype, but a few paratypes (KU 328735, 328736, TNHC 61990, 61994) possess darker throats, owing to aggregation of black blotches as in the holotype. Summaries of univariate morphological variation in the series are presented in Table 2. Females (PNM 9907 ��� 0 9, 9914) are considerably larger and more robust (Table 2), and the single juvenile (KU 339544; SVL 24.3 mm) was more brightly patterned in life, with bolder and contrasting light tan versus black blotches. Distribution. Platymantis quezoni is known only from the type locality but may be more wide spread in scattered karst outcrops of northern portions of the Bicol Peninsula (Restificar et al. 2006; MGB 2010), especially on limestone karst formations in the nearby Bondoc Peninsul, Published as part of Brown, Rafe M., Layola, Louise Abigail De, Ii, Antonio Lorenzo, Diesmos, Mae Lowe L. & Diesmos, Arvin C., 2015, A new species of limestone karst inhabiting forest frog, genus Platymantis (Amphibia: Anura: Ceratobatrachidae: subgenus Lupacolus) from southern Luzon Island, Philippines, pp. 191-210 in Zootaxa 4048 (2) on pages 194-204, DOI: 10.11646/zootaxa.4048.2.3, http://zenodo.org/record/239892, {"references":["Brown, W. C., Brown, R. M. & Alcala, A. C. (1997 a) Species of the hazelae group of Platymantis from the Philippines, with descriptions of two new species. Proceedings of the California Academy of Sciences, 49, 405 - 421.","Brown W. C., Alcala, A. C. & Diesmos A. C. (1999) Four new species of the genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines. Proceedings of the California Academy of Sciences, 51, 449 - 460.","Brown, W. C., Alcala, A. C., Diesmos, A. C. & Alcala, E. (1997 c) 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. & Alcala, A. C. (1970) 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.","Siler, C. D., Alcala, A. C., Diesmos, A. C. & Brown, R. M. (2009) A new species of limestone-forest frog, genus Platymantis (Amphibia: Anura: Ceratobatrachidae) from eastern Samar Island, Philippines. Herpetologica, 65, 92 - 104. http: // dx. doi. org / 10.1655 / 08 - 040 R. 1","Brown, R. M & Gonzales, J. C. (2007) A new forest frog of the genus Platymantis (Amphibia; Anura; Ranidae) from the Bicol Peninsula of Luzon Island, Philippines. Copeia, 2007, 251 - 266. http: // dx. doi. org / 10.1643 / 0045 - 8511 (2007) 7 [251: ANFFOT] 2.0. CO; 2","Brown, W. C. & Alcala, A. C. (1982 a) A new cave Platymantis (Amphibia: Ranidae) from the Philippine Islands. Proceedings of the Biological Society of Washington, 95, 386 - 391.","Restificar, S. D. F., Day, M. J. & Urich, P. B. (2006) Protection of karst in the Philippines. Acta Carsologica, 35, 121 - 130. http: // dx. doi. org / 10.3986 / ac. v 35 i 1.248","Mines and Geosciences Bureau (MGB) (2010) Geology of the Philippines. Mines and Geosciences Bureau, & Department of the Environment and Natural Resources, Quezon City, 311 pp.","Brown, W. C., Alcala, A. C. & Diesmos, A. C. (1997 b) A new species of the genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines. Proceedings of the Biological Society of Washington, 110, 18 - 23."]}

Brown, Rafe M., Layola, Louise Abigail De, Ii, Antonio Lorenzo, Diesmos, Mae Lowe L., Diesmos, Arvin C. (2015): A new species of limestone karst inhabiting forest frog, genus Platymantis (Amphibia: Anura: Ceratobatrachidae: subgenus Lupacolus) from southern Luzon Island, Philippines. Zootaxa 4048 (2): 191-210, DOI: http://dx.doi.org/10.11646/zootaxa.4048.2.3

ALCALUS GEN. NOV. Type species Micrixalus mariae Inger 1954. Diagnosis Members of the genus Alcalus can be distinguished from many members of the clade Anurajen (species of the genera Cornufer and Platymantis) by having (1) an intermediate body size (Al. baluensis: males 20– 25 mm Snout-Vent Length (SVL), females 26–31; Al. mariae: males 32–37 mm SVL, females 35–43); (2) a broad head (vs. slender to moderately broad); (3) a coarsely textured, shagreened, or ‘wrinkled’ skin appearance in all species (vs. smooth, tuberculate, or with longitudinal dorsolateral dermal ridges); (4) widely expanded, terminally squared, spatulate toe discs (vs. non- or minimally expanded, terminally rounded); (5) semi-aquatic microhabitat preferences (vs. preferences for terrestrial or arboreal microhabitats in most species); and by the (6) presence of nuptial pads in males (vs. absence); (7) absence of vocal sacs (vs. presence of median subgular vocal sacs); (8) absence of supernumerary tubercles on hand (vs. presence in most species); (9) presence of elongate subarticular tubercles (vs. presence, round); (10) absence of outer metatarsal tubercles on plantar surface of feet (vs. presence in most species); and (11) presence of extensive, usually full, interdigital webbing of the feet (vs. absence). Phylogenetic definition Alcalus (NCN) is a maximum crown-clade name that refers to the crown clade (C) originating in the last common ancestor of Al. mariae and all extant species that share a more recent common ancestor with Al. mariae than with Cornufer vitiensis or Platymantis corrugatus. It can also be conceived of as the largest crown clade containing Al. mariae, but not Co. vitiensis or Pl. corrugatus. Content Southeast Asian (Sunda Shelf and Palawan Island) species formerly placed in Ingerana (Table 3): Al. mariae, Al. baluensis, Al. rajae, and presumably Al. sariba (Shelford, 1905), which was not sampled (Table 3). Comment It is not surprising that the montane, semi-aquatic, Southeast Asian island archipelago species formerly referred to Ingerana comprise a monophyletic group, unrelated to the ecologically dissimilar and biogeographically disjunct mainland species of Ingerana (as presently understood, from Andaman Islands, Bhutan, China, north eastern India, Myanmar, and Nepal). Erection of a new genus to accommodate these taxa is undertaken here with reference to the phylogenetic placement of the type species of ‘true’ Ingerana (I. tenasserimensis), which in our phylogeny is more closely related (but with weak support) to the Dicroglossidae than to the Ceratobatrachidae (Fig. 2). The placement of Alcalus as the sister group of the clade Anurajen (containing genera Platymantis and Cornufer) has been confirmed elsewhere (Bossuyt et al., 2006; Frost et al., 2006; Wiens et al., 2009; Pyron & Wiens, 2011), although taxon sampling was not as extensive. The phylogenetic relationships and possible additional generic subdivision of the nonceratobatrachid (perhaps dicroglossid) species referred to Ingerana remain unstudied. Etymology A masculine noun honouring our long-term collaborator, friend, and mentor Angel C. Alcala for his numerous contributions to the systematics, ecology, conservation, and developmental biology of Southeast Asian amphibians. Suggested common name: Alcala’s dwarf mountain frogs.

Brown, Rafe M., Siler, Cameron D., Richards, Stephen J., Diesmos, Arvin C., Cannatella, David C. (2015): Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 174 (1): 130-168, DOI: 10.1111/zoj.12232, URL: http://dx.doi.org/10.1111/zoj.12232

CERATOBATRACHINAE BOULENGER, 1884 Diagnosis Other species of Ceratobatrachinae differ from Alcalus by having (1) a relatively narrow head (vs. wide); and (2) smooth skin, with or without dermal tubercles and/ or dermal ridges (vs. coarsely textured, shagreened, or ‘wrinkled’ in appearance); (3) rounded terminal toe discs (vs. spatulate toe discs with squarish terminal shape); (4) absence of nuptial pads in males (vs. presence); (5) presence of a medial subgular vocal sac in most species (vs. absence); (6) presence of palmar supernumerary tubercles (vs. absence); and (7) presence of rounded subarticular tubercles and outer metatarsal tubercles on plantar surface (vs. presence, elongate). Finally, most species of Ceratobatrachinae (except Cornufer guppyi, species of the subgenus Potamorana, and Platymantis levigatus) differ from Alcalus by the absence of interdigital webbing on the feet (vs. presence); and by having terrestrial or arboreal microhabitat preferences (vs. semiaquatic). Morphological synapomorphies for Ceratobatrachinae have not been identified. Content The genera Cornufer and Platymantis (see below). Comment We have not converted Ceratobatrachinae (Clade D) to a clade name, but have instead coined a new unranked clade name, Anurajen. If Ceratobatrachinae were converted to a clade name, and if the family Ceratobatrachidae (Node B) were re-ranked as Ceratobatrachinae in the future, then the phylogenetic name Ceratobatrachinae and the ranked name Ceratobatrachinae would refer to different clades, which would cause confusion. Therefore, we define a new clade name below denoting the same clade as the subfamily Ceratobatrachinae., Published as part of Brown, Rafe M., Siler, Cameron D., Richards, Stephen J., Diesmos, Arvin C. & Cannatella, David C., 2015, Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae), pp. 130-168 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 174 (1) on page 143, DOI: 10.1111/zoj.12232, http://zenodo.org/record/5337577, {"references":["Boulenger GA. 1884. Diagnoses of new reptiles and batrachians from the Solomon Islands, collected and presented to the British Museum by H. B. Guppy, Esq., M. B., H. M. S. ' Lark'. Proceedings of the Zoological Society of London 1884: 210 - 213."]}

ALCALINAE SUBFAM. NOV. Type genus Alcalus (see account below). Diagnosis The diagnosis for Alcalinae is the same as for the genus Alcalus, below. Phylogenetic definition We have not defined Alcalinae as a phylogenetic name because it would be redundant with Alcalus; it adds no new information about phylogenetic relationships. However, we name the ranked subfamily Alcalinae, even though it is also redundant in content with Alcalus, to provide a coordinate name for its sister-taxon Ceratobatrachinae. Content One genus, Alcalus, which includes the species Al. mariae, Al. baluensis, and Al. rajae. We anticipate that Ingerana sariba eventually will be transferred to the new genus as well. Etymology See Etymology section for the genus Alcalus below.

SUBGENUS PLATYMANTIS GÜNTHER, 1858 Diagnosis The subgenus Platymantis (currently a single recognized species, Pl. corrugatus) differs from other species of Platymantis by having (1) elongate longitudinal dermal ridges along the dorsal body surfaces (vs. dorsum smooth or tuberculate); (2) distinctive ‘quaaack’ advertisement calls (vs. frequency sweeps, pure tones, or complex calls); and (3) distinctive dark lateral head coloration (of varying shades; vs. lateral head pigment undifferentiated from surrounding coloration). The diagnostic dark lateral head coloration forms a dark ‘facemask’ that we consider a synapomorphy of this clade. Additionally, members of this subgenus can be diagnosed from species of arboreal variable Philippine forest frogs of the genera Lahatnanguri (Platymantis banahao, Pl. cornutus, and Pl. insulatus), and all members of the subgenera Tahananpuno and Tirahanulap, by the absence of expanded digital tips of fingers and toes (vs. presence of some degree of terminal digital expansion of fingers and toes), and by having a terrestrial microhabitat preference (normally calling beneath leaf litter) and a crepuscular (vs. nocturnal) calling activity pattern (Table 3). Content The allopatric populations of (1) the Luzon and West Visayan faunal regions, (2) the Camiguin Norte lineage, (3) the populations from the Mindanao faunal region islands, and (4) the Mindoro Island populations, all currently referred to Pl. corrugatus (Table 3). The subgenus Platymantis is equivalent in content to the unranked taxon Tagomukhus. Conversion of the name Platymantis would result in two different clades, ranked as genus and subgenus, with the same name, Platymantis. Therefore we define a NCN, denoting the same clade as subgenus Platymantis., Published as part of Brown, Rafe M., Siler, Cameron D., Richards, Stephen J., Diesmos, Arvin C. & Cannatella, David C., 2015, Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae), pp. 130-168 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 174 (1) on page 144, DOI: 10.1111/zoj.12232, http://zenodo.org/record/5337577, {"references":["Gunther ACLG. 1858. Neue Batrachier in der Sammlung des britischen Museums. Archiv fur Naturgeschichte, Berlin 24: 319 - 328."]}

Platymantis citrinospilus sp. nov. Figs. 2���4 Holotype. South Australia Museum (SAMA) R 64758 (SJR Field No. 10860), adult male, collected by S. J. Richards at Tompoi Camp, 1700 m above sea level (05�� 20.623 'S, 151 �� 18.873 'E; WGS- 84), Nakanai Mountains, East New Britain Province, Papua New Guinea, 23 April 2009. Paratypes. SAMA R 64756, R 64757 (SJR Field Nos. 10831, 10832), and Papua New Guinea National Museum (PNGNM) 24042 (SJR Field No. 10811) three adult males, same locality data as holotype, SAMA R64756, 64757 collected 22 April 2009, and PNGNM 24042 collected 21 April 2009. Etymology. The specific epithet, a masculine name, is derived from the Greek adjective kitrinos (yellow) and the Greek noun spilos (spot or stain), in reference to the distinctive bright yellow flank areolations of all known specimens (Fig. 2). Diagnosis. Platymantis citrinospilus is assigned to the genus Platymantis on the basis of established diagnostic characters for the genus (Small body size, lack of interdigital webbing, presence of distinct subarticular tubercles, expanded terminal phalanges of the hand and foot, and nuptial pads absent; Brown 1952; Gorham 1965; Menzies 2006) and it is distinguished from congeners by (1) body size (29.5���32.2 mm in four males), (2) widely expanded terminal disks of the fingers and toes (Figs. 2, 3), (3) skin of dorsum smooth to finely granular with enlarged dermal, fleshy tubercles, (4) ventral surface of subarticular tubercles on fingers, toes, hands and feet uniformly low and flattened, (5) fingers and toes without lateral dermal flange, (6) interdigital webbing absent, (7) tibio-tarsal articulation with prominent, ornate white dermal tubercles, (8) flank with prominent yellow areolations, (9) intraocular region with low but distinct mid-sagittal crest, and (10) unique advertisement call consisting a relatively slow series of dull, high-frequency notes (resembling the sound produced by slowly striking together two pieces of wood). Comparisons.���The new species is phenotypically most similar to (and likely most closely related to) a pair of other Nakanai Mountains shrub frogs, P. macrosceles and P. mamusiorum. Platymantis citrinospilus differs from P. macrosceles by the presence of brick reddish-brown ground coloration (Fig. 2) with bright yellow flank areolations (versus homogenous bright green body coloration), by the presence of low supra-ocular and tarsal tubercles (versus presence of highly enlarged, white and prominent tubercles; Fig. 2), by the presence of a conspicuous mid-saggital crest between the eyes (versus absence), by its mottled bronze-brown iris (versus light lime green iris), and by its higher frequency, slowly-pulsed advertisement call (versus lower frequency, rapidly pulsed call). Platymantis citrinospilus is also a high elevation species, known only from 1700 m above sea level in the Nakanai Mountains while Platymantis macrosceles has only been recorded at elevations between 800 and 900 m above sea level (Foufopoulos & Brown, 2004; Foufopoulos & Richards, 2007); neither species has been recorded at intervening elevations despite multiple surveys (Foufopoulos & Brown, 2004; Foufopoulos & Richards, 2007). Platymantis citrinospilus differs from syntopic P. mamusiroum by the presence of fleshy dermal dorsal tubercles (versus absence; interocular region flat), by its more pointed (versus rounded) snout and sharp (versus sloping) canthus, by its brick reddish-brown dorsal ground color and lateral yellow flank areolations (versus bright dreen dorsum with fine brown reticumum), by the presence (versus absence) of supra-ocular and tarsal tubercles, and by its pulsed advertisement call (versus a stridulated series of ���croaks��� or ���crunches��� in P. mamusiorum). Of the other tree-dwelling shrub frogs of New Britain, Platymantis citrinospilus (SVL 29.5���32.2) differs from the larger P. nakanaiorum (SVL 35.8 ���38.0) and P. nexipus (SVL 39.3���43.7) and the much smaller P. caesiops (SVL 18.9���22.9) by body size, and more extensive dorsal, supra-ocular, and extensive (versus minimal) tarsal tuberculation, and pulsed (versus click and buzzes) advertisement call (Foufopoulos & Brown, 2004; Brown et al., 2006 a; Kraus & Allison, 2009) (Table 1). Platymantis citrinospilus can easily be distinguished from all terrestrial species of Platymantis on New Britain (P. adiastolus, P. akarithymus, P. boulengeri, P. bufonulus, P. gillardi, P. magnus, P. mimicus, P. schmidti, and P. sulcatus) by the presence of widely expanded digital disks of fingers and toes (versus finger and toe termini non-expanded to slightly expanded); additional morphological and acoustic differences are summarized by Brown and Tyler (1968), Brown and Menzies (1979), Brown et al., (2006 a,b), and Kraus and Allison (2007). Description of holotype ���A mature male, in excellent condition; habitus slender; head slightly distinct, equal in dorsal aspect to width of body, HL 36.6 % SVL; HL 100 % HW; snout moderate, terminating in bluntly round point in lateral and doral aspects, protruding only sigthly beyond lower jaw; eyes protrude only slightly beyond silhouette of head in dorsal aspect, and moderately beyond dorsal surface of head in lateral aspect; labial region slightly flared, not extending beyond eyes in dorsal aspect; interorbital region with a moderately prominent, longitudinally-oriented crest; ED 120 % IOD; pupil horizontally ovoid; canthus rostralis markedly medially bowed; loreal region very concave; ED 61.0% SNL; narial openings slightly laterally protuberant; eye-narial distance 5 times the distance from nostril to tip of snout; internarial region flat; tympanum distinct, TD 56 % ED; dorsal edge of tympanic annulus partially concealed by supratympanic fold, the latter extending from dorsoposterior edge of eye, over dorsal edge of tympanum, and terminating at supra-axillary (post-rictal) region; two or three conical postrictal tubercles present between posteroventral edge of tympanum and forearm insertion (one much larger than others); tongue triangular, with shallow posterior notch and narrow anterior attachment; choanae round, minute, at anterolateral edge of palate, separated by a distance 6���8 times their diameter, not obscured by palatal shelf; dentigerous process of vomer ovoid; vomerine teeth minute, translucent, numbering two or three; dentigerous process anterolaterally angled, with closest (posterior) points separated by a distance 1.5 times the diameter of one choana, their most distant (anterior) ends separated by a distance equal to three or four times diameter of choanae; openings to vocal sac minute slits, at the level of the angle of the jaw. Skin of dorsal surfaces of body, head, and limbs finely granular, with ornamentation in the form of moderate fleshy conical dermal tubercles (lacking obvious keratin) in supra-orbital, post-orbital, and post-rictal regions (Fig. 2); dermal tubercles of lateral body surfaces and dorsal limb surfaces low and irregularly scattered; two infralabial tubercles barely visible in ventral and lateral aspect at the outer (posterior) margins of the jaw; low fleshy tubercular ridges present on lateral edges of radii and sacral region; irregular row of prominent tubercles extends along the posteromedial (postaxial) edge of tarsal segment of the hind limb (Fig. 3), clustered most prominently and densely clustered at tibio���tarsal articulation; ventral surfaces of trunk, head, and throat, slightly glandular, becoming more coarsely glandular in posterior quarter of trunk, groin, and medial half of the humeri. Hand length 65.6 % PL; fingers (Fig. 4) wide and round in cross-section, (lateral dermal flanges absent); terminal disks barely expanded (Finger I) to widely expanded (> 2 times the width of penultimate phalanges in Fingers II���IV), with circum-marginal folds encircling distal edges of disks; supra-articular folds indistinct, more evident above penultimate and ultimate phalangeal articulation of inner two fingers; decreasing finger length III, IV, II, I; subarticular tubercles low, mostly flat on ventral surfaces, not pointed, the most prominent angled slightly anteriorly towards tip of fingers; one subarticular tubercle under fingers I���II, two tubercles under fingers III���IV; supernumerary tubercles flat and somewhat indistinct, present at the base of fingers II���IV; palmar surfaces basal to supernumerary tubercles nearly smooth; thenar (inner metacarpal), medial palmar and outer metacarpal tubercles moderate, flat on ventral surfaces, edges irregular but distinct, unpigmented; thenar tubercle irregularly ovoid, situated on medial edge of finger I; medial palmar (inner metacarpal) tubercle ovoid on left, elongate on right, only moderately enlarged, with slightly distinct, raised medial edge, approximately the same size as thenar tubercle; outer metcarpal tubercles small, ovoid, slightly more than half the size inner metacarpal tubercle and distinct (separated from medial metacarpal tubercle); nuptial pads absent, forearm musculature not well developed or hypertrophied. Species: P. citrinospilus P. macrosceles P. mamusiorum P. nakanaiorum P. nexipus P. caesiops (4) (4) (6) (4) (16) (2) SVL (mm) 29.5���32.2 29.4���31.8 27.4���30.7 35.8 ���38.0 39.3���43.7 18.9���22.9 Hindlimbs long; TBL 58.5 % SVL, PL 83.2 % TBL; skin of dorsal hind limb surfaces smooth; tarsus smooth, with prominent posteriorly oriented conical tubercles; low conical tubercles distributed along postaxial dermal tarsal flange and at heel; tubercles most prominent and clustered around tibio-tarsal articulation; terminal toe disks non-expanded (Toe V), narrowly expanded ( 2 times the width of penultimate phanges; Toes II, III), with circum-marginal grooves and supra-articular cutaneous folds; plantar surface of feet (Fig. 4) smooth, devoid of supernumerary tubercles, with well-developed, prominently rounded (not pointed) subarticular tubercles; subarticular tubercles with flattened ventral surfaces, three under Toe IV, two under Toes III and V, and one under Toes I and II; decreasing toe length (longest to shortest when adpressed) IV, III, V, II, I; outer metatarsal tubercle small, round, pointed; separated from inner metatarsal tubercle by distance three times width of outer; inner metatarsal tubercle low, flat, moderate, oblong, with slightly sharp medial edge, four to five times size of outer metatarsal tubercle; toes without interdigital webbing. Cloacal region coarsely glandular, with small supra-cloacal dermal flap; low, fleshy cluster of enlarged white tubercular swellings present ventrolaterally on each side of vent. Measurements of holotype ���SVL 32.2; ED 3.9; TD 2.2; HL 12.8; SNL 6.1; END 3.9; IOD 2.6; HW 11.9; FL 16.0; TBL 17.5; TSL 9.7; PL 15.3; ML 10.3; FA 5.1; Toe 4 L 9.5; Fin 1 L 4.3; Fin 3 L 3.8; Fin 3 DW 2.4; Fin 3 PPW 0.8; Toe 4 DW 1.1; Toe 4 PPW 0.8. Coloration of holotype in preservative ���Dorsal surfaces of body, head and limbs dark brick reddish-brown (ground coloration of dorsal and lateral surfaces of head distinctly darker), with scattered diffuse steel gray-green pigmentation clustered most densely on snout, eyelids, forearms, midbody and sacral regions, knees, and tibial segments of hindlimbs; congregation of this pigmentation feature forms a fine vertebral line from tip of snout to vent; grey-green pigment densely clustered along outer margins of eyelids, continuous with lateral snout pigmentation and forming a prominent palpebral���canthal ���stripe;��� dark dorsal color continues laterally on to brick reddish-brown flanks, with prominent series of bold, highly distinct, yellow spots and blotches running in a continuous series from inguinal region, across all ventrolateral surfaces, to supra-articular region above forelimb insertion (terminating below tympanum); a similar, bold marking is angled obliquely across the tympanum (anteriodorsally to posteroventrally); labial region similarly bright pale yellow, nearly devoid of any dark pigment; dorsal surfaces of forelimbs light brick reddish-brown, with less dense steel gray-green pigment than dorsal surfaces of body, lacking transverse bars; dorsal surfaces of hands and fingers light reddish-brown, with differentiated absence of brown (resulting in pale cream color in unpigmanted areas) on articulations of penultimate���ultimate phalanges; dorsal surfaces of finger disks light cream with brown blotches (Fingers III and IV) or bright cream (Fingers I and II); dorsal surfaces of hindlimbs orange-brown on femur, steel gray-green on tibia, and orangish-brown on tarsus, transverse bars absent; concealed surfaces on posterior thighs orangish-brown with large bright pale yellow spots, similar to those on flanks; dorsal surfaces of toes light brown with slightly darker terminal disks on outer toes (IV and V) and lighter cream on inner toes (I���III); ventral surfaces orangish brown with distinct, bold pale yellow spots, wrapping on to ventrum from lateral surfaces; underside of throat homogenous orangish-brown contrasting with bright pale yellow lower lip; ventral surfaces of forelimbs reddishbrown, with irregular bold pale yellow markings on upper arms, some steel gray-green pigment wrapping ventrally from lateral surfaces, bright pale yellow forearm tubercles, and pale cream subarticular tubercles and ventral surfaces of terminal finger disks; ventral surfaces of glandular skin of trunk orangish, fading to pale gray midventrally; ventral surfaces of hindlimbs slightly darker orangish brown with a few bold pale yellow spots on femoral segments; ventral surfaces of foot homogenous reddish-brown with pale yellow to cream subarticular tubercles and gray terminal disks. Variation ���Our small type series exhibits some notable color variation but the types are generally phenotypically similar to one another. The holotype (SAMA R 64758) exhibits markedly more steel gray-green pigmentation than the three paratypes, although PNGNM 24042 has this same pigment on its limbs and head, and all specimens have this color feature on eyelids and lateral surfaces of limbs. The paratypes all exhibit dorsal coloration darker than that of the holotype. SAMA R 64756 and R 64757 have notably lighter, homogenous orangish-brown dorsal coloration. All specimens possess the faintly lighter, thin vertebral line from snout to vent (least evident in SAMA R 64756). No specimens have transverse bars on limbs, and all paratypes have nearly homogenous orangish-brown dorsal limb color. The bold lateral pale yellow flank areolations are present in all specimens but vary in size and density (Fig. 2). The holotype (SAMA R 64758) and paratype PNGNM 24042 exhibit a series of large areolations that connect into a near-continuous wavy line, whereas SAMA R 64756 possesses nearly discrete, unconnected flank spots. SAMA R 64757 has discrete spots on the left flank but fewer, partialy connected spots on the right. In SAMR R 64756 and 64757, flank areolations are bordered by thin black lines. Bold yellow flank spot series continue to the infratympanic region (above forearm insertion) and extend onto the tympanum in the holotype (SAMA R 64758) and paratype PNGNM 24042 while in paratypes SAMR R 64756 and 64757, distinct pale yellow spots extend only to the ventral edge of the tympanum. In these latter specimens, tiny pale yellow spots are also present at the center of the tympanum. All specimens exhibit bright, pale yellow upper and lower labial coloration which is interrupted by orangish-brown pigment in SAMR R 64756 and 64757. Ventral coloration is nearly invariant and similar to that of the holotype except that darkened pigment on the throat (congregation of dark pigment marking the positions of the vocal sacs) is more evident in SAMA R 64756 and 64757, and in SAMA R 64758 more bright pale yellow spots extend from flanks on to the ventral surface of the trunk. Variation in bright pale yellow spot configurations on the posterior surfaces of the thighs and the cloacal region is as follows: in the holotype (SAMA R 64758) spots on the distal portions of the posterior surfaces of the thighs are distinct and separate, and gradually fuse together to form a continuous supra-cloacal marking; in PNGNM 24042, a bright pale yellow supra-cloacal bar is present and distal thigh spot are absent; and in SAMR R 64756 and 64757 only a few small, intermittently fused, bright pale yellow spots are present in the cloacal region. Ventral surfaces of the hands and feet are largely invariant, although subarticular tubercles and terminal disks of SAMA R 64756 and 64757 are cream to light gray whereas these same structures are pale yellow in paratype PNGNM 24042 and the holotype (SAMA R 64758). Coloration in life ���Based on color images of all specimens in life (images of the holotype, SAMA R 64758, and paratypes PNGNM 24042 and SAMA R 64756 are included in Fig. 2). Dorsal ground coloration dark brick reddish-brown with extensive (SAMA R 64758), limited (PNGNM 24042) or nearly absent suffusion of light green pigment on dorsal surfaces of head, trunk and limbs (Fig. 2). When juxtaposed with white dermal tubercles on dorsal surfaces, overall pale green lichenatious color results on dorsum (strongest in holotype SAMA R 64758). All specimens have a thin light cream to white vertebral stripe, a bright immaculate white (SAMA R 64756) to yellow (SAMA R 64758, PNGNM 24042) labial and postrictal region, with (SAMA R 64758, PNGNM 24042) or without (SAMA R 64756, PNGNM 24042) intrusion of bright postrictal coloration on to ventral edge of the tympanum. Lateral head color similar to dorsum; iris pale brown above and below the pupil. Flank surfaces and concealed posterior surfaces of the thighs with bold, bright yellow areolations varying from numerous (10���12) small but distinct yellow spots (SAMA R 10831) to four or five larger blotches with irregular borders. Dorsal limb surfaces were colored as body, with indistinct slightly darker reddish-brown transverse bars present (PNGNM 24042) or absent (remainder). The dorsal surfaces of the digits were brick reddish-brown; expanded digital disks white. Ventrum was pale pink, with dark red throat, and reddish color wrapping slightly on to ventrum from lateral surfaces (Fig. 3 D). In life the ventral body surfaces of SAMA R 64756 and PNGNM 24042 were somewhat translucent, with darker and lighter organs visible though the skin. The ventral surfaces of limbs were dark red with bold yellow spots; the ventral surfaces of all specimens��� hands and feet were dark red with pale yellow (PNGNMR 24042) to cream subarticular tubercles and terminal ventral surfaces of digital disks (remainder). Advertisement call and acoustic comparisons to P. macrosceles ���The following call descriptions are based on two recording segments approximately of two minutes per species. Calls of P. citrinospilus (both individuals vouchered, SAMA R 64758 and R 64756) and P. macrosceles (UWZM 23721, another individual recorded but not collected) were all recorded at ambient temperatures of 16.9 ���18.0��C. Platymantis citrinospilus calls (defined here as sounds produced per single expiration) consist of a relatively slow series of dull, medium-frequency notes sounding, Published as part of Brown, Rafe M., Richards, Stephen J. & Broadhead, Taylor S., 2013, A new shrub frog in the genus Platymantis (Ceratobatrachidae) from the Nakanai Mountains of eastern New Britain Island, Bismarck Archipelago, pp. 31-45 in Zootaxa 3710 (1) on pages 33-40, DOI: 10.11646/zootaxa.3710.1.2, http://zenodo.org/record/284660

Brown, Rafe M., Richards, Stephen J., Broadhead, Taylor S. (2013): A new shrub frog in the genus Platymantis (Ceratobatrachidae) from the Nakanai Mountains of eastern New Britain Island, Bismarck Archipelago. Zootaxa 3710 (1): 31-45, DOI: http://dx.doi.org/10.11646/zootaxa.3710.1.2

Platymantis biak sp. nov. Figs. 2–4 Holotype. PNM 9679 (ACD Field No. 4543; KU 319973), adult male, collected on limestone outcrop in secondary-growth forest (20:00 hr) on 17 January 2009 in Barangay Biak na Bato, Municipality of San Miguel and Doña Remedios Trinidad, Bulacan Province, Luzon Island, Philippines (N 15.1084, E 121.0724; 190 m elevation), by ACD Paratopotypes. Twenty-three males (KU 319977 –78, 319981–82, 319987, PNM 9680, 9681, 9682 –83, 9684– 9697), eight females (KU 319974 –75, 319979, PNM 9674 –75, 9676, 9677 – 78), and seven juveniles (KU 319976, 319980, 319983, PNM 9698–9701) collected on karst formations in secondary-growth forest (18:00– 23:00), January–March and November 2009, in Barangay Biak na Bato, Municipality of San Miguel and Doña Remedios Trinidad, Bulacan Province, Luzon Island, Philippines; collected by ACD, CDS, CWL, and. MLD. Diagnosis. We assign the new species to the genus Platymantis on the basis of (1) its moderate body size, (2) absence of interdigital webbing, (3) the presence of a forked omosternum, (4) directly developing reproductive mode, and (5) terrestrial microhabitat preference (Brown and Alcala 1980; Brown et al., 1997; Brown, 2004). Platymantis biak can be distinguished from other Philippine congeners by the following combination of characters: body size 32.3–39.9 mm SVL for males, 37.4–42.4 mm SVL for females; digital disc of Finger III 1.9–2.2 × width of penultimate phalanx in males and 1.4–2.4 in females (Fig. 3); Finger I slightly longer than Finger II; finger discs larger than toe discs; smooth skin; spotted flanks absent; supratympanic fold smooth, less protuberant; tympanum completely visible; dorsolateral stripes absent; dorsal folds, ridges, dermal tubercles absent; and a preference for terrestrial microhabitat in karst forest. Mensural and diagnostic differences are provided in Tables 1 and 2. Description of holotype. A mature male; habitus robust; head broader than body, head length 35.5 % snout–vent length, 90.4 % head width; snout protruding moderately beyond lower jaw, snout tip comes to moderately sharp point in dorsal and lateral aspects; eyes moderate and protruding anterolaterally, eye diameter 12.7 % snout–vent length, 35.9 % head length; lips slightly flared and swollen (Fig. 2); interorbital region slightly convex; eye diameter 80.9 % snout length, 134.2 % eye–narial distance; pupil horizontally elliptical; canthus rostralis anteriorly concave in dorsal aspect; loreal region moderately concave, sloping ventrolaterally to labium; nostrils oriented posterolaterally; eye–narial distance 60.3 % snout length; internarial region slightly concave; tympanic annulus distinct 56.8 % eye diameter; supratympanic fold slightly protuberant, extending from posterior corner of eye across dorsal margin of tympanum to supraaxillary region. Tongue elongate, wider posteriorly, with deep posterior notch and narrow anterior attachment; choanae situated at anterolateral edge of palate, oval, separated by distance two times greater than diameter of single choana; dentigerous process of vomer distinct, with no teeth visible on process; dentigerous process angled slightly anterolaterally, approximately at 20–30 º incline with closest (posterior) points separated by distance equal to 2.0 × diameter of single choana, their most distant (anterior) ends separated by distance equal to 3.0 × diameter of single choana. Hand length 54.6 % foot length; tibia length 113.0% foot length, 54.3 % snout–vent length; fingers slender, long; terminal discs moderately expanded (Fig. 3 A) with pronounced distal, circum-marginal groove; dorsal surfaces of terminal phalanges with small, cutaneous supra-articular flaps between ultimate and penultimate phalanges; relative lengths of fingers II FIGURE. 2. Photographs in life of Platymantis biak paratopotypes exhibiting (A) typical limestone-forest frog mottled color pattern, PNM 9678, female, SVL = 40.7 mm, and (B, C) perching behavior observed at type locality, PNM 9674, female, SVL = 40.8 mm. Photographs by A. Diesmos and C. Linkem. Skin of dorsal and ventral surfaces of head, trunk, and limbs smooth. Terminal discs of toes slightly expanded, with circum-marginal grooves; supra-articular cutaneous flaps above ultimate-penultimate phalangeal articulations; plantar surface of foot smooth, with well-developed, prominently rounded subarticular tubercles (Fig. 3 B); supernumerary tubercles absent; relative lengths of toes I Measurements of holotype (in mm). Snout–vent length 39.9; head length 14.2; head width 15.7; snout length 6.3; upper arm length 9.0; forearm length 9.3; tibia length 21.7; femur length 20.4; tarsus length 9.4; foot length 19.2; hand length 10.5; Finger I length 7.6; Finger III length 7.6; Toe IV length 10.4; eye–narial distance 3.8; snout–narial distance 2.0; interorbital distance 3.3; internarial distance 3.2; eye–tympanum distance 0.8; eye diameter 5.1; horizontal tympanic annulus diameter 2.9; Finger I disc width 1.2; Finger III disc width 1.8; Toe IV disc width 1.2; Finger III penultimate phalange width 0.6; Toe IV penultimate phalange width 0.5. FIGURE. 3. Ventral views of hand (A) and foot (B) of female paratopotype of Platymantis biak (KU 319974), hand (C) and foot (D) of male Platymantis dorsalis (KU 300356), and hand (E) and foot (F) of male Platymantis guentheri (KU 306324). Scale bar = 2 mm. Coloration of holotype in preservative. The dominant dorsal color pattern on the head and body consists of dark mottled blotches of black interspersed with blotches of dark shades of brownish green. The dorsal surfaces of the limbs are banded with alternating black and dark green bars, separated by thin tan bands. An interorbital bar is present consisting of a thin, irregular, light-brown line. The head and snout region are darker than the dorsum, and nearly uniform black. A black bar extends across the dorsal half of the canthus rostralis to the anterior edge of eye. The tympanum is medium brownish-green with a dark blotch extending posterodorsally from the center. The temporal region is predominantly black with several small blotches of dark green. Lip coloration consists of alternating dark green and black vertical labial bars, bordered anteriorly by black lower edge of canthal and subocular regions. The transverse bars on the limbs have irregular widths, with an overall lighter coloration when compared to the dorsum due to the greater proportion of skin covered by tan and dark green bands. Arm banding patterns coalesce into mottled patterns on the elbows. Four black bars are present on the forearms, thighs, and tibia, with three bars present on the tarsus. Anterior flank coloration is predominantly black, progressively fading to lighter grayish-tan with dark brown blotches and spots, blending into a black, dark brown, and tan mottled pattern posteriorly. The dorsal surfaces of the hands are light brown, with alternating bands of light brown and grayish-tan on Fingers I–III, and dark brown and grayish-tan on Finger IV. The dorsal surfaces of the feet are colored with alternating bands of light brown and grayish-tan. The throat and chest are marbled light cream with several small tan speckles, while the periphery of the throat and lower lips are darker, with the anterior edge of the lower lip possessing an irregular outline of dark green blotches. The belly is homogeneous light tan bordered by light cream spots laterally. Ventral coloration on the arms consists of light tan coloration present on the upper arm and interior portions of the forearm, and dark green coloration present on the outer portions of the forearm and hand. Ventral surfaces of the thighs uniformly mottled cream anteriorly and light green-brown posteriorly. This coloration blends gradually down the legs into a homogenous dark brown color pattern on the tarsus and foot, similar to the dark coloration observed on the forearms. The subarticular tubercles of the hands and feet are light cream, and the iris is dark blue-gray around the pupil. FIGURE. 4. Photographs in preservative of Platymantis biak holotype (PNM 9679) in dorsal and lateral views of head. Scale bar = 5 mm. Table 1. continued. Coloration 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; lower surfaces of supratympanic fold black; black canthal blotch located just anterior to eye; black labial blotch located below eye, anterior to tympanic annulus; dorsal surfaces of limbs with alternating dark brown and green transverse bars, each bordered by thin cream transverse bars; dorsal surface of body with several large black blotches, irregularly distributed; area of black blotches and cream speckles on anterior portion of flanks; pupil bordered by copper ciliary ring of iris, sharply blending into predominant mottled gold and brown color pattern of iris. biak sp. nov. bayani insulatus spelaeus paengi (12m, 4 f) (9m, 9 f) (8m, 6 f) (4m, 3 f) (15m) SVL (m) 32.3–39.9 34.2–39.1 34.3 –38.0 37.1–44.3 27.7–34.3 SVL (f) 37.4–42.4 44.3–49.8 40.2–42.9 53.5– 57.5 N/A Range Central Luzon Samar Island Gigantes Islands SW Negros NW Panay Island Island Island Species Group guentheri guentheri guentheri dorsalis dorsalis Digital disks Moderately Widely Widely Non-expanded Non-expanded expanded expanded expanded Dorsal skin rugosity – – to granular – +, – + Dorsal dermal tubercles – +, sparse – +, small + Flanks Blotches anteriorly Spotted Spotted Immaculate Spotted Variation. Dominant color pattern consistently mottled in all observed specimens. Variation exists in the w-shaped pattern of pigmentation on the dorsal surface of the body in line with the forearms. Some individuals possess a sharply defined w-shaped pattern (KU 319976, 319980, 319982 –83, 319987, PNM 9684 –86, 9688–89, 9691, 9693, 9695, 9697, 9700), while the pattern on many other specimens is either obscured by the background body coloration or is vaguely defined (KU 319973 –75, 319977–79, 319981, PNM 9698 –99, 9680–83, 9674–76, 9687, 9690, 9692, 9694, 9696, 9677 –78, 9701, 9704 – 14). The outer metacarpal tubercle was fully divided in most individuals with the exception of a few individuals showing a single, large, connected outer metacarpal tubercle (KU 319974, 319980, 319981, 319983, 319987). Females are considerably larger and more robust. Summaries of univariate morphological variation in the series are presented in Table 3. Distribution. Platymantis biak is known only from the type locality (Fig. 1). Ecology and natural history. Platymantis biak occurs in secondary-growth forest on karst substrate. Both males and females were observed perched on top of limestone rocks within large rock formations in the forest and around the entrances to cave systems. Additionally, males were observed up to 5 meters inside caves, some perched above the ground within depressions and holes in the karst rock walls (Fig. 2 B, C). No specimens were observed perching in trees or shrubs, in contrast to the common behavior of most species in the P. guentheri Group (Brown and Gonzales, 2007). Some individuals of Platymantis biak were well camouflaged against the limestone (much like P. bayani, P. insulatus, P. speleaus, and P. paengi; Fig. 2 B, C), and field workers found it difficult to locate and capture individuals against the limestone substrate. This was especially true for many of the females. However, males were often more conspicuous while calling atop rocky outcrops. When disturbed, the new species immediately jumped back into the rocks and escaped into limestone crevices. No egg masses or juveniles were found; other species of Platymantis have direct development (Brown and Alcala, 1982 a, 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 (Brown and Alcala, 2000). We assume that this species also breeds in limestone crevices and small caves (Brown and Alcala, 2000; Brown et al., 2003). Sympatric anuran species include Rhinella marina (introduced), Hoplobatrachus rugulosus (introduced), Kaloula picta, K. pulchra (introduced), Limnonectes macrocephalus, L. woodworthi, Occidozyga laevis, Platymantis cf. mimulus, Polypedates leucomystax, Rana erythraea (introduced), and R. similis. Etymology. We derive the specific epithet from the Tagalog term biak, meaning "crevice" or "crack," in reference to the preferred limestone karst habitat at the type locality, Biak na Bato (meaning "cracked rock" or "fragmented stone"), Bulacan Province. Comparisons. The new species is assignable to the P. g u e n t h e r i Group (P. banahao, P. bayani, P. cornutus, P. diesmosi, P. guentheri, P. insulatus, P. luzonensis, P. negrosensis, and P. rabori) based on the following combination of characters: digital discs moderately expanded and larger in fingers than in toes, proximal portions of digits with narrow dermal folds in cross section, and subarticular tubercles large and highly protuberant (Brown et al., 1997 c; Fig. 3). The new species is morphologically most similar to Platymantis bayani and Platymantis insulatus, the other two members of the P. g u e n t h e r i Group with similar morphology and preference for karst forest microhabitats (Tables 1 and 2). Platymantis biak is distinguished from P. bayani and P. insulatus by the presence of moderately (vs. widely) expanded digital discs and the absence of fully spotted flanks (vs. spotted). From Platymantis bayani, the new species differs by its smaller female body size (maximum SVL 42.4 mm for P. biak vs. 49.8 mm for P. bayani), smaller eye diameter (maximum ED 5.2 mm for P. b i a k vs. 6.8 mm for P. bayani), and the absence of granular skin (vs. absence or presence) and dorsal dermal tubercles (vs. presence). The new species further differs from Platymantis insulatus by the presence of well-defined limb banding patterns (vs. absence or presence and poorly defined). Additional characters for distinguishing the new species from all species of the P. guentheri Group are summarized in Tables 1 and 2. From all members of the P. d o r s a l i s 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. t a y l o r i), Platymantis biak is distinguished by the presence (vs. absence) of moderately expanded digital discs (Fig. 3) and larger finger discs than toe discs. Platymantis biak is further distinguished from all members of the P. dorsalis Group, except P. levigata by the absence of dorsal folds and ridges, dermal tubercles, and rugose skin (vs. presence of one, two, or all characters), 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. t a y l o r i, 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. t a y l o r i, 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 slightly longer than the second; narrow dermal phalanges on the digits; protuberant (vs. flattened) subarticular tubercles, and a preferred microhabitat of terrestrial substrates in limestone forest (vs. arboreal habitat, and/or shrub-layer vegetation).

Brown, Rafe M., Richards, Stephen J. (2008): Two new frogs of the genus Platymantis (Anura: Ceratobatrachidae) from the Isabel Island group, Solomon Islands. Zootaxa 1888: 47-68, DOI: 10.5281/zenodo.184280

Platymantis desticans sp. nov. Figs. 2, 3 A, 4 A Holotype. SAMA R 56849, adult male, collected by S. Richards at Barora Faa Island, (off the western tip of Isabel Isl.), 0���10 m above sea level (07�� 32.065 'S, 158 �� 21.005 'E), Isabel Province, Solomon Islands, 10 May 2000. Paratypes. SAMA R 56850 ��� 52, three adult males, same data as holotype except SAMA R 56852 collected on 11 May 2000. Referred Specimens. Solomon Island, Choiseul Province, Choiseul Island, Pavora River: two specimens deposited in the Australian Museum corresponding to ABTC 50323 and 50385 tissue vouchers. Etymology. The specific epithet is an adjective derived from the Latin verb destico meaning to squeak, and is used in reference to this species��� distinctive advertisement call. Diagnosis. Platymantis desticans is distinguished from congeners by a combination of (1) body size (36.9 ���38.0 mm for four adult males), (2) narrowly expanded terminal disks of the fingers and toes (Fig. 2), (3) skin of dorsum shagreened with fine dermal asperities and scattered irregular tubercular ridges, (4) subarticular tubercles of fingers, toes, hands and feet very prominent and raised, (5) plantar surface of pes smooth between subarticular tubercles, (6) digits lacking all vestiges of lateral flange and interdigital webbing, and (7) unique advertisement call. The new species is morphologically most similar to P. w e b e r i but differs from this species by differences in body size and proportions (most notably hind limb length, reflected in TBL/SVL ratio, Table 1), the presence (vs. absence) of dermal asperities on the dorsum, and the presence of short, irregular tubercular ridges (vs. highly prominent, elongate, longitudinally oriented and medially bowed dermal ridges and folds), by a smooth (vs granular) plantar surface of the pes between subarticular tubercles (Fig. 2), by dark brown (vs. light cream) ventral coloration, and by its highly distinctive advertisement call (Fig. 5). The new species is also phenotypically similar to P. myersi (from Bougainville Island) but differs from that species by the presence of dermal asperities on the dorsum (vs asperities absent, dorsum smooth or weakly tuberculate posteriorly), a short, rounded (vs long, pointed) snout, the complete absence of interdigital webbing (vs minute web present between toes in P. myersi), thighs smooth (vs strongly granular), plantar surface of pes smooth between subarticular tubercles (vs strongly granular) and an apparently uniform color pattern (vs color pattern variable; Brown, 1949, 1952). Description of holotype. A mature male, in excellent condition; habitus slender; head slightly distinct, only slightly wider in dorsal aspect than body, length 45 % of SVL; head length 114 % of head width; snout short, its tip bluntly rounded, protruding slightly beyond lower jaw, rounded in lateral aspect; eyes protrude laterally beyond silhouette of head in dorsal aspect, and moderately beyond dorsal surface of head in lateral aspect; labial region rounded and smooth, only slightly flared, lips not swollen, not extending past eyes when viewed in dorsal aspect; interorbital region flat; eye diameter 162 % of interorbital distance; pupil horizontally ovoid; canthus rostralis barely medially bowed; loreal region slightly concave; eye diameter 62.9 % of snout length; nostrils not laterally protuberant; eye-narial distance 6 times the distance from nostril to tip of snout; internarial region flat; tympanum distinct, its diameter 72 % of eye diameter; dorsal edge of tympanic annulus partially concealed, overlapped by supratympanic fold; the latter extending from dorsoposterior corner of eye, along dorsal edge of tympanum, and terminating nearly at supraaxillary region; cluster of five strongly conical post-rictal tubercles present between posteroventral edge of tympanum and forearm insertion; tongue ovoid, with deep posterior notch and narrow anterior attachment; choanae round, minute, at anterolateral edge of palate, separated by a distance five to eight times greater than their diameter, nearly obscured by palatal shelf; dentigerous process of vomer elongate; vomerine teeth tiny, translucent, numbering six to eight; dentigerous process very slightly angled anterolaterally, with closest (posterior) points separated by a distance two times the diameter of one choana, their most distant (anterior) ends separated by a distance equal to five or six times diameter of choanae; openings to vocal sac moderate slits, at the level of the angle of the jaw. Skin of dorsal surfaces of body, head, and limbs generally roughened by dense congregation of dermal asperities (Figs. 3���4; giving the skin the texture of fine sandpaper), and with short irregular tubercular ridges scattered at various angles across dorsum; suprascapular dermal folds absent; dorsal surfaces of limbs textured as body but tubercular ridges absent; ventral surfaces of trunk, head, throat, and limbs smooth; manus length 53.9 % of foot length; digits of manus narrow (Fig 2 A) and round in cross-section; lateral edges of digits lack fleshy dermal flanges; terminal disks slightly expanded and protuberant, with short circum-marginal folds on distal tips of digits; minute supra-articular flaps present above penultimate-ultimate phalangeal articulation; decreasing digital length III, I, IV, II; subarticular tubercles extremely prominent and protuberant, rounded on ventral surfaces; one subarticular tubercle on digits I���II, two tubercles under digits III���IV; supernumerary tubercles very prominent and rounded, present at the base of all digits; surface of palmar surfaces basal to supernumary tubercles entirely smooth; thenar (inner metacarpal), medial palmar and outer metacarpal tubercles prominent, highly convex; thenal tubercle greatly enlarged and protuberant, subtriangular and nearly twice the size of (and separated completely from) medial and outer metacarpals; medial palmar ovoid and larger than elongate, outer metacarpal slender; medial and outer metacarpal tubercles not separated; nuptial pads absent, forearm musculature not hypertrophied. Hindlimbs short; tibia length 48.9 % of snout-vent length, foot length 96.6 % of tibia length; tarsus smooth, lacking folds, flaps, or tubercles; terminal disks of toes moderately expanded, protuberant, and ovoid; lateral dermal flanges absent along distal phalanges; circum-marginal grooves limited to distal ends of digits; supraarticular cutaneous folds slight; plantar surfaces of pes smooth (Fig. 2 B), with prominently protuberant subarticular tubercles but lacking supernumerary tubercles or texture on plantar surfaces; subarticular tubercles numbering three under Toe IV, two under Toes III and V, and one each under Toes I and II; decreasing digit length IV, III, V, II, I; outer metatarsal tubercle prominent and circular, pointed; inner metatarsal tubercle moderate, low, elongate, with a slightly sharp venterolateral edge; digits of pes completely free of web; supracloacal tubercles and dermal flaps absent. Species P. parilis P. guppyi P. neckeri (4 m) (18 m) (12 m) SVL (mm) 39.5���41.4 44.5���63.4 38.6���47.4 Range Barora Faa, Isabel, Choiseul Throughout the Solomons Bougainville Measurements of holotype. SVL 38.0; ED 5.0; TD 3.6; HL 17.1; SNL 7.9; IOD 3.1; HW 15.0; FL 18.3; TBL 18.6; TSL 9.9; PL 18.0; ML 9.7; FA 8.2; Toe 4 L 13.8; Fin 1 L 5.7; Fin 3 L 7.2; Fin 3 DW 0.9; Fin 3 PPW 0.5; Toe 4 DW 0.9; Toe 4 PPW 0.6. Color of holotype in preservative. In preservative, the holotype exhibits a nearly homogeneous dark brown dorsal coloration, with fainter medium brown blotches irregularly scattered across the trunk and a prominent light parietal spot on the back of the head posterior to the orbits; dark dorsal coloration blends laterally into light and dark brown marbled flanks with irregular and indistinct tan blotches; dorsal and lateral surfaces of snout colored as body, tympanum slightly lighter, and distinct cream labial bars present; post-rictal region heterogeneous, with distinct cream-tipped post-rictal tubercles; dorsal surfaces of limbs lighter brown than body, with darker (similar to trunk coloration) transverse bars (three on forearm, three on humerus, three on tibia, two on tarsus); dorsal surfaces of hands, feet, and digits similarly colored brown with dark spots and white bars on phalangeal articulations; venter pale medium brown with distinct dark brown mottling; underside of throat heavily suffused with brown with dark brown labial bars wrapping on to ventral surfaces of lower lips; ventral surfaces of hands dark brown with prominent white subarticular tubercles; ventral surfaces of limbs very dark brown with only faint vestiges of lighter coloration; ventral surfaces of hands and feet solid dark brown with light gray, velvety digital disks. Va r i a t i o n. There is very little color variation evident in the type series. One specimen (SAMA R 56851) is noticeably lighter in dorsal coloration, has a lighter venter, and pale brown spots on the throat. The remaining paratypes agree almost perfectly with the holotype in every respect. Color in life. Based on color images of holotype in life, before preservation (Fig. 3). Dorsal surfaces homogeneous dark chocolate brown with distinct white parietal spot. Labial bars alternated black and white, in contrast to the brown coloration on lateral surfaces of head and snout. The tympanum coloration was bluegray and black and lower portions of the flanks were blue-gray, wrapping on to the venter. The iris was gold above and blue below the pupil and post-rictal tubercles were distinctly white-tipped. The dorsal surfaces of the digits were brown with black transverse bars and white on spots on digital articulations. The throat and venter were dark chocolate brown, fading to tan by the groin. Ventral surfaces of limbs were dark reddish brown. Advertisement call. The advertisement call of P. desticans is markedly distinct from that of any known species of Platymantis in both temporal and spectral characteristics (Fig. 6). The following description is based on two vouchered recordings from SAMA R 56849 (holotype) and 56850 (paratype). The call (defined throughout as vocalizations produced during a single expiration) is a high-frequency tonal note (sounding to the human ear like a shrill ���squeak���), and is repeated in a slow train, characterized with variable intercall interval at the beginning (or ���warm-up��� period) of a calling bout. As the animal begins to call steadily, calling repetition rate structure becomes steadily discernable. During peak calling activity, the call is delivered in groups of paired calls, separated from one another by a nearly invariant intercall interval and with each pair of calls separated by a lengthy period of silence. Individual call group repetition rate (i.e., rate of delivery of paired call groups) was 0.07 calls/s for the holotype (SAMA R 56849) and 0.08 calls/s for the paratype (SAMA R 56850). Call repetition rate (not taking into account paired call group structure; = [total number of calls ��� 1]/time from beginning of first call to beginning of last) was 0.13 SAMA R 56849 and 0.16 for SAMA R 56850. For paired calls, mean intercall interval was 585.1���1242 ms (= 975.8 �� 27 SD; n= 4) for SAMA R 56849 and 627.3���1411 ms (= 945.3 �� 41 SD; n= 6) for SAMA R 56850. Duration of silence between call groups (i.e., long interval between paired calls) was 9.8��� 18.6 s (= 13.0 �� 3.1 SD; n= 4) for SAMA R 56849 and 21���32 s (= 24.7 �� 6.5 SD; n= 6) for SAMA R 56850. Call structure of P. d e s t i c a n s is complex and invariant. The shrill, high frequency single-note squeak call of P. desticans has an elaborate internal temporal and spectral structure (Fig 6 B, C). Each note contains two distinct components: an amplitude modulated and frequency modulated first component and then a second, frequency modulated, constant amplitude component. There are pronounced differences in call energy contained in the different components although peak amplitude of both components is nearly equivalent (Fig. 5 B). Although the audiospectrogram reveals cryptic stucture within the call of P. desticans, the impression on the human ear is a single, slightly wavering, shrill squeak. The first intracall component of P. desticans is a brief frequency arc. The dominant frequency for the arc component began at 2.1���2.3 kHz, climbed to peak at 3.2���3.4 kHz, and then declined to 2.1���2.3 kHz in SAMA R 56849. In SAMA R 56850, the frequency arc peaked slightly higher at 3.4���3.5 kHz. The shape of this arc was nearly symmetrical in SAMA R 56849 (Fig. 6 B) but contained a slight ascending prefix tail (not shown) in SAMA R 56850. The arc component varied in duration by 26.0��� 29.1 ms (= 27.6 �� 2.1 SD; n= 5) in SAMA R 56849 and 29.6���32.8 ms (= 30.9 �� 2.7 SD; n= 8) in SAMA R 56850. In the arc component of all calls, the vast majority of calling energy is packaged into the dominant frequency, described above. However in some calls, a lower frequency (fundamental) frequency component is discernable in addition to one or two higher frequency harmonics (multiples of the fundamental). In our recording of SAMA R 56849, the fundamental frequency begins at 1.08 kHz, climbs to 1.8 kHz, then declines again to 1.1 kHz. Above the dominant frequency are two harmonics that peak at 4.9 and 6.7 kHz, respectively. In the recording of SAMA R 56850, the fundamental frequency arc climbs from 1.0 to 1.9, before declining again to 1.1 kHz. In all available recording segments, the arc prefix component of the call of new species exhibits an inverse relationship between call frequency and call amplitude (Fig. 6 B). Call energy is most intense when frequency of the arc component is at its lowest, this amplitude falls as frequency increases, the n climbs again as frequency declines The second component of the P. desticans call is a gradual frequency sweep. In SAMA R 56849, frequency of the second component declines from 2.9 to 2.7 kHz and in SAMA R 56850 the frequency sweep component declined from 2.9 to 2.7 kHz. The second component varied in length from 66.7 to 80.3 ms (= 75.4 �� 9.3 SD; n= 15) in calls of SAMA R 56849 and from 71.2 to 84.3 ms (= 77.1 �� 11.1 SD; n= 10) in SAMA R 56850. Ecology and Natural History. Platymantis desticans occurs either at extremely low densities or is naturally very inconspicuous, as no additional specimens were seen or heard despite intensive searches during the survey. Males were detected predominantly by their soft squeaky calls, uttered from exposed positions on the forest floor. Two males were calling from muddy banks near streams and one from the base of a fallen log. The call of this species was extremely difficult to hear at distances greater than about 10 m. In contrast Platymantis weberi occurred in the same habitats at extremely high densities and their calls dominated the acoustic environment at this site. Other species occurring in sympatry with the new species were: Ceratobatrachus guentheri, Discodeles guppyi, Palmatorappia solomonis, Platymantis aculeodactylus, P. guppyi, P. neckeri, P. solomonis and P. w e b e r i. Distribution. Confirmed to date at two localities: Barora Faa Island, Isabel Province and Pavora River, Choiseul Island, Choiseul Province., Published as part of Brown, Rafe M. & Richards, Stephen J., 2008, Two new frogs of the genus Platymantis (Anura: Ceratobatrachidae) from the Isabel Island group, Solomon Islands, pp. 47-68 in Zootaxa 1888 on pages 49-56, DOI: 10.5281/zenodo.184280, {"references":["Brown, W. C. (1949) A new frog of the genus Platymantis from the Solomon Islands. American Museum Novitates, 1387, 1 - 4.","Brown, W. C. (1952) The amphibians of the Solomon Islands. Bulletin of the Museum of Comparartive Zoology, Harvard, 107, 1 - 64."]}

Platymantis parilis sp. nov. Figs 3 B, 4 B, 6, 7 Holotype. SAMA R 56911, adult male, collected by S. Richards near Kolopakisa Village, ~ 0���10 m above sea level (07�� 36.008 'S, 158 �� 39.092 'E), north-western Isabel Isl., Isabel Province, Solomon Islands, on 16 May 2000. Paratypes. SAMA R 56908 ��� 10, three males, same data as holotype except SAMA R 56910 collected on 17 May 2000. Etymology. The specific epithet is chosen from the Latin parilis, meaning equivalent, like, or similar, in reference to the morphological similarity between the new species and the acoustically and genetically distinct Platymantis neckeri. Diagnosis. Platymantis parilis is distinguished from congeners by (1) body size (39.5���41.5 mm for four adult males), (2) widely expanded terminal disks of the fingers and toes (Fig. 6), (3) skin of dorsum entirely smooth or with only a few low tubercular ridges in the scapular region, (4) subarticular tubercles of fingers, toes, hands and feet low and flattened on their ventral surfaces, (5) fingers and toes with lateral dermal flange, (6) toes with basal vestiges of interdigital webbing, and (7) unique advertisement call. Platymantis parilis is morphologically indistinguishable from P. neckeri but cannot be confused with any other species of Platymantis. Aside from genetic data indicating that P. parilis and P. neckeri consistently form monophyletic clusters of moderately genetically distinct haplotypes (3.7���4.2 % uncorrected percent divergence in the 16 S ribosomal RNA mitochondrial gene between four samples of P. parilis and 16 samples of P. neckeri; RMB, unpublished data), the new species can be readily distinguished by its unique advertisement call (Figs. 7, 8). Description of holotype. A mature male, in excellent condition; habitus slender; head slightly distinct, only slightly wider in dorsal aspect than body, length 40.2 % of SVL; head length 102 % of head width; snout moderately long, its tip sharply pointed, protruding well beyond lower jaw, pointed in lateral aspect; eyes protrude laterally only slightly beyond silhouette of head in dorsal aspect, and well beyond dorsal surface of head in lateral aspect; labial region flat and slanted, lips flared, protuberant, extending well beyond eyes in dorsal aspect; interorbital region flat; eye diameter 118 % of interorbital distance; pupil horizontally ovoid, nearly subcircular; canthus rostralis nearly straight (slightly medially bowed); loreal region very slightly concave, continuous with flat slanting upper lip; eye diameter 50.9 % of snout length; nostrils laterally protuberant; eyenarial distance 4���5 times the distance from nostril to tip of snout; internarial region flat; tympanum distinct, its diameter 63.7 % of eye diameter; dorsal edge of tympanic annulus in contact with, but not concealed by supratympanic fold, the latter extending from dorsoposterior corner of eye, along dorsal edge of tympanum, and terminating nearly at supra-axillary region; a single weakly conical post-rictal tubercle present between posteroventral edge of tympanum and forearm insertion; tongue triangular, with deep posterior notch and narrow anterior attachment; choanae round, minute, at anterolateral edge of palate, separated by a distance 5���6 times their diameter, not obscured by palatal shelf; dentigerous process of vomer ovoid; vomerine teeth tiny, translucent, numbering 4 or 5; dentigerous process sharply anterolaterally angled, with closest (posterior) points separated by a distance two times the diameter of one choana, their most distant (anterior) ends separated by a distance equal to 4���5 times diameter of choanae; openings to vocal sac moderate slits, at the level of the angle of the jaw. Skin of dorsal surfaces of body, head, and limbs smooth; a few low and indistinct short tubercular ridges in scapular region; suprascapular dermal folds and dorsolateral folds absent; dorsal surfaces of limbs textured as body; tubercles absent; ventral surfaces of head, throat and limbs smooth; trunk weakly granular; groin slightly more granular; manus length 58.9 % of foot length; digits of manus (Fig. 6 A) wide and ovoid in crosssection, owing to lateral dermal flange bordering fingers; lateral edges of all digits with wide fleshy dermal flanges; terminal disks widely expanded, exceeding twice the width of penultimate phalanges, with circummarginal folds wrapping around distal portions of digits; supraarticlar flaps present above penultimate-ultimate phalangeal articulation; decreasing digital length III, IV, II, I; subarticular tubercles low, not pointed, angled anteriorly towards tip of digits, flattened on ventral surfaces; one subarticular tubercle under digits I��� II, two tubercles under digits III���IV; supernumerary tubercles flat and somewhat indistinct, present at the base of all digits; palmar surfaces basal to supernumerary tubercles entirely smooth; thenar (inner metacarpal), medial palmar and outer metacarpal tubercles flat, edges indistinct; thenal tubercle greatly moderate and elongate, situated on medial edge of digit I, connected to medial dermal flange; medial palmar (inner metacarpal) tubercle greatly enlarged, subtriangular, twice the size of thenar tubercle; outer metcarpal tubercles small, subcircular, half the size of thenar tubercle, not separated from medial metacarpal tubercle; nuptial pads absent, forearm musculature not hypertrophied. Hindlimbs short; tibia length 50.1 % of snout-vent length, foot length 95.6 % of tibia length; tarsus without minute dermal flaps but with a few very low tubercles, continuous in line with postaxial dermal flange along Toe V and outer edge of pes (terminating after wrapping around ankle); small tubercles present on tibio-tarsal articulation; digits of toes widely expanded (Fig. 6 B), twice the width of penultimate phanges; circummarginal grooves wrapping around distal ends of digits; supraarticular cutaneous folds present above penultimateultimate phalangeal articulation; plantar surfaces of pes smooth, with low subarticular tubercles but lacking supernumerary tubercles or texture on plantar surfaces; subarticular tubercles with flattened ventral surfaces, numbering three under Toe IV, two under Toes III and V, and one each under Toes I and II; decreasing digit length IV, III, V, II, I; outer metatarsal tubercle low, flattened, indistinct and circular; inner metatarsal tubercle moderate, low, elongate, twice the size of outer metatarsal tubercle, with only a slightly sharpened venterolateral edge; digits of pes basally webed; web reaching first subarticular tubecles of digits all digits, except extending half way between the first and second subarticular tubercle on the outer edge of Toe III; supracloacaal tubercles and dermal flaps absent. Measurements of holotype. SVL 39.5; ED 4.1; TD 2.7; HL 16.7; SNL 7.7; IOD 3.3; HW 15.5; FL 20.3; TBL 20.6; TSL 910; PL 19.3; ML 11.5; FA 8.8; Toe 4 L 13.3; Fin 1 L 4.3; Fin 3 L 8.1; Fin 3 DW 1.9; Fin 3 PPW 1.1; Toe 4 DW 1.8; Toe 4 PPW 1.0. Color of holotype in preservative. Dorsal surfaces dark brown with pale yellow streaks and blotches irregularly scattered across the trunk and head; eyelids and interorbital region yellowish-tan; dark dorsal coloration fades laterally into light yellowish-tan flanks with irregular brown blotches; dorsal and lateral surfaces of snout colored as body, with brown background coloration and yellowish-tan loreal region, tympanum, and labial bars; tympanum and post-rictal tubercle streaked with pale yellow; snout brown with yellow nares and tip of snout; dorsal surfaces of forelimbs pale yellow, with blotches of gray-brown on wrists, lacking transverse bars; dorsal surfaces of hindlimbs tan with gray-brown blotches, transverse bars absent; concealed surfaces on posterior thighs orange; dorsal surfaces of hands and fingers pale yellow with few brown flecks, lacking differentiated coloration on phalangeal articulations; dorsal surfaces of toes medium gray-brown with black to gray terminal disks; venter pale yellowish cream; underside of throat heavily flecked with slight brown and with medium brown labial blotches wrapping on to the lower lips; ventral surfaces of forelimbs yellowish, with darker brown blotches on outer edges, wrapping around from lateral surfaces; ventral surfaces of hindlimbs slightly darker, orangeish-cream; ventral surfaces of hands pale cream with cream subarticular tubercles and terminal disks; ventral surfaces of pes medium tan and subarticular tubercles and terminal disks gray. Va r i a t i o n. The paratypes all exhibit dorsal coloration darker than that of the holotype. The three specimens are nearly homogeneously dark brown, with slightly lighter interorbital regions and snouts. All have indistinct transverse bars on limbs, numbering two on forearm, two on tibial segment, and a single bar on the tarsus. In these more darkly-colored specimens, there is a distinct demarcation on the flanks between dark dorsal coloration and yellowish-tan ventral color. In one paratype (SAMA R 56908) this stratification occurs more dorsally, and the flanks are predominantly yellow with distinct dark blotches; in SAMA R 56906, the dark-light demarcation occurs ventrally, and the flanks are dark brown with bright yellow blotches. In paratype SAMA R 56909 the loreal region is distinctly lighter than the rest of the lateral surfaces of the head and in all paratypes the post-rictal tubercles are bright white. In all specimens, the cartilaginous protrusion at the tip of the snout is pale cream. In the three paratypes, the throats are distinctly darker than the holotype, owing to infusion of small, scattered flecks; in these same specimens, ventral trunk coloration is pale cream with brown flecks and ventral surfaces of hindlimbs are darker brown. In all paratypes, the ventral surfaces of the hands are darker than that of the holotype, with dark brown to gray palmar coloration and light cream subarticular tubercles and terminal disks; ventral surfaces of the feet are dark brown with dark gray tubercles and terminal disks. Color in life. Based on color images of paratype SAMA R 56908 in life, before preservation (Fig. 3 B). Dorsal surfaces homogeneous light brown with light interorbital bar and slightly lighter snout. Lateral head color similar to dorsum, but with darker streak across tympanum and a few dark blotches on the upper lips. The iris was silvery above and blue below the pupil and post-rictal tubercles were distinctly white-tipped. The upper arm was distinctly purple in life and remaining dorsal surfaces of the limbs were colored as body, but with indistinct slightly darker brown transverse bars. The dorsal surfaces of the digits were tan with dark brown. Advertisement call of P. parilis (and comparison to the call of P. n e c k e r i ). The following call descriptions are based on two approximately 1 min recording segments per species (all recorded by SJR, within 2 ��C of each other; P. parilis SAMA R 56911 and P. neckeri SAMA R 56792). Platymantis parilis calls (defined here as sounds produced per single expiration) with a simple, fast train of low frequency (dominant frequency = 1.3���1.5 kHz), dull sounding, dual frequency component notes, lacking any tonal elements (Fig. 7). The impression on the human ear is the sound of wood blocks striking together very rapidly with a constant rate of note delivery. Calls of individual males of P. parilis contained 11���14 (mean 12.6 �� 2.7 SD) notes per call. Each note contains two distinct frequency components (Fig. 7 A, B) with a simple, amplitude modulated structure within each call (Fig. 7 B���F). The lower dominant frequency component of each note peaked at 1.3 kHz and 1.5 kHz. The higher frequency component varied from 2.7 kHz for one specimen and 3.0 kHz for the other. In all calls, the fundamental (lower) frequency component was the dominant (= emphasized) frequency at the start of each call. Towards the end of many calls, the two frequency components were approximately equivalent in relative intensity, suggesting a slight change in call energy from the fundamental to the first multiple of the fundamental (the higher frequency component of the call). However, the dominant call frequency never shifted markedly away from the fundamental (Fig. 7 B, D). The calls of P. parilis appeared stereotyped and nearly invariant. Some variation was detected in dominant frequency between males (1.3���1.5 kHz) but we found little or no within-male or within-call frequency modulation, except for the aforementioned slight shifts in call energy towards the higher frequency component. Although P. neckeri also produces an amplitude-modulated call (Fig. 8), the structure and overall impression on the human ear is unlike that of P. p a r i l i s. The call of P. n e c k e r i sounds to humans like a rapid, clanging ���chi-keng���..chi-keng ���chi-keng ���chi-keng���chi-keng���chi-keng���chikeng!��� with an increasing rate of note delivery, often terminating at a rapid burst of notes. Platymantis neckeri has a lower mean note repetition rate (4.7���9.6 ���notes/s) than P. parilis (11.8���12.1 notes/s) and produces fewer notes per call (5���7) than P. parilis (11���14). The call of P. neckeri exhibits a progressively decreasing internote interval across the call, whereas P. parilis has a uniform internote interval throughout the entire call. Individual note structure between the two species was also quite different. In contrast to Platymantis parilis, which exhibits 11���14 invariant dual-frequency note structure across the call, the call of P. n e c k e r i typically delivers 12 notes of a mean peak dominant frequency of 3.2���3.4 kHz that alternate back and forth between two structurally different syllables (Fig 8 B). Menzies (1982 b) described the calls as an invariant 6 -note call, with each note divided into two or three syllables (depending on the presence of the brief suffix added to the tonal note in some calls). The first note is a dual-frequency pulse with two distinct frequency components. This is followed by a second note consisting of a simple tone or shallow declining tonal frequency sweep. This two-note sequence is repeated 5���7 times (Menzies, 1982 b, reported this repeated sequence as invariably six). Unlike P. parilis, which exhibits a two-frequency note structure wherein the fundamental was usually the dominant frequency (but occasionally calls in which the two frequencies were equivalent), P. n e c k e r i invariably possessed calls in which the dominant frequency corresponds to the higher frequency component of the call. Additionally, unlike P. p a r i l i s, which produces a harmonically simple call, the calls of P. neckeri have a rich harmonic structure, with clear multiples of the dominant frequency occurring at 6.7���6.9 and 10.1���10.3 kHz (not shown). Ecology and Natural History. The new species called exclusively from elevated sites (typically leaves), normally between 2 and 6 meters above the ground in dense rainforest. At least one specimen appeared to be calling from within an epiphyte; others were perched on leaves in dense foliage. Other species occurring in sympatry with P. parilis included Ceratobatrachus guentheri, Discodeles guppyi, Palmatorappia solomonis, Platymantis aculeodactylus, P. guppyi, P. neckeri, P. solomonis and P. w e b e r i. In sympatry, the advertisement calls of P. parilis and P. neckeri were remarkably invariant and consistently different. Our impression at the type locality was that P. p a r i l i s is much more abundant than P. neckeri. Distribution. Known only from the vicinity of Kolopakisa village, north-western Isabel Island., Published as part of Brown, Rafe M. & Richards, Stephen J., 2008, Two new frogs of the genus Platymantis (Anura: Ceratobatrachidae) from the Isabel Island group, Solomon Islands, pp. 47-68 in Zootaxa 1888 on pages 56-62, DOI: 10.5281/zenodo.184280, {"references":["Menzies, J. I. (1982 b) The voices of some male Platymantis species of the New Guinea region. British Journal of Herpetology, 6, 1 - 5."]}

Platymantis adiastola, new species (Figs 4���6) Holotype: SAMA R 61906, adult male, collected by S. Richards at Wanui Camp, Wanui River Valley (near Wide Bay), 310 m above sea level (05 21.638 ���S, 152 �� 05.266���E), New Britain Isl., East New Britain Province, Papua New Guinea, 20 March 2000. Paratypes: SAMA R 57014 ��� 15, R 57040 ��� 41, same data as holotype except collected 13 March 2000; R 60257 ���8, 15 March 2000; R60259, 18 March 2000; SAMA R61907, 19 March; SAMA R 61908, UPNG 9974 -6, 20 March 2000; SAMA R61909, 22 March 2000. Etymology: The specific epithet is chosen from the Greek adiastolos, meaning confused, mixed up, or not separated, in reference to past confusion surrounding the taxonomic status of the new species. Diagnosis: Platymantis adiastola is distinguished from congeners by (1) body size (39.5���43.9 for 11 males; 71.7 for one female), (2) narrowly expanded terminal disks of the fingers and toes, (3) skin of dorsum with scattered longitudinal tubercular ridges, including two long (4���6 mm) medially bowed suprascapular dermal ridges, (4) upper eyelids devoid of conical tubercles, (5) subarticular tubercles of fingers, toes, hands and feet prominent and raised, (6) digits lacking all vestiges of lateral flange and interdigital webbing, and (7) unique advertisement call. The new species is morphologically nearly identical to P. schmidti but differs from this species by slight differences in body size and proportions (Table 1), an apparent tendency towards a more elevated calling perch preference, and a possible greater extent of sexual size dimorphism (P. adiastola male/female SVL = 0.57; P. schmidti = 0.69). The main distinguishing feature between these two morphologically cryptic forms is the male advertisement call. Platymantis adiastola produces a call composed of a series of shrill, high frequency (mean peak dominant frequency 3.0��� 3.2 kHz), complex, tonal notes whereas P. schmidti calls with a simple train of twin-pulse lower frequency (2.4���2.6 kHz) notes lacking tonal elements. Platymantis adiastola has a lower mean note repetition rate (4.3 notes/s) than P. s c h m i d t i (12.8 notes/s) and produces fewer mean notes per call (mean = 16.6) than P. s c h m i d t i (mean = 23.9). Finally, the call of P. adiastola exhibits a progressively decreasing internote interval across the call, whereas P. s c h m i d t i has a uniform internote interval throughout the entire call. Description of holotype: A mature, undissected male specimen, in excellent condition; habitus moderately stocky; head distinct, slightly wider in dorsal aspect than body, length 42.6 % of SVL; head length 114.2 % of head width; snout short, its tip bluntly rounded, protruding slightly beyond lower jaw; snout posteroventrally slanted in lateral aspect; eyes protrude laterally beyond silhouette of head in dorsal aspect, and strongly beyond dorsal surface of head in lateral aspect; labial region flat and smooth, not flared, lips not swollen, not extending past eyes when viewed in dorsal aspect; interorbital region flat; eye diameter 143.4 % interorbital distance; pupil horizontally ovoid; canthus rostralis slightly concave; loreal region gently sloping, slightly concave; eye diameter 73.0% of snout length, 126.6 % of eye���narial distance; nostrils slightly laterally protuberant; eye narial distance much longer than distance from nostril to tip of snout; internarial region flat; tympanum distinct, its diameter 56.9 % of eye diameter; dorsal edge of tympanic annulus in contact with and slightly overlapped by supratympanic fold, the latter extending from dorsoposterior corner of eye, along dorsal edge of tympanum, and Platymantis adiastola Platymantis schmidti Platymantis schmidti n. sp. (original collections) (recent collections) terminating nearly at supraaxillary region; tongue subcircular, with deep posterior notch and narrow anterior attachment; choanae round, minute, at anterolateral edge of palate, widely separated by a distance eight to ten times greater than their diameter, not obscured by palatal shelf; dentigerous process of vomer oblong; vomerine teeth tiny, translucent, numbering four to six; dentigerous process very slightly angled anterolaterally, with closest (posterior) points separated by a distance two times the diameter of one choana, their most distant (anterior) ends separated by distance equal to three times diameter of choanae; openings to vocal sac short slits, slightly posterior to rictus; prominent postrictal tubercle cluster present at posteroventral edge of tympanum. Skin of dorsal surfaces of body, head, and limbs generally smooth, except for fine longitudinal ridges of variable length on dorsum; a faint pair of medially bowed suprascapular dermal folds present; several other short glandular folds present along dorsolateral edge of trunk and in temporal region; ventral surfaces of trunk, head, throat, and limbs smooth; manus length 45.9 % of foot length; digits of manus narrow in appearance and round in cross-section; lateral edges of digits without fleshy dermal flanges; terminal disks moderately expanded and protuberant, with short circumarginal folds on distal tips of digits; minute supraarticlar flaps present above penultimate-ultimate phalangeal articulation; decreasing digital length III, I, II=IV; subarticular tubercles extremely prominent and protuberant, rounded on ventral surfaces; one subarticular tubercle on digits I���II, two tubercles under digits III���IV; supernumerary tubercles very prominent and rounded, present at the base of all digits; minute palmar tubercles preset basal to supernumary tubercles; thenar (inner metacarpal), medial palmar and outer metacarpal tubercle prominent, ovoid, highly convex, and nearly equal in size; slightly raised dermal tubercles present on ventral surfaces of wrists; nuptial pads absent, forearm musculature not hypertrophied. Hindlimbs moderately well developed; foot length 91.9 % of tibia length; tibia length 60.3 % of snout-vent length; tarsus smooth, lacking folds, flaps, or tubercles; digits of toes moderately expanded and ovoid; slight, narrow lateral dermal flanges present along distal phalanges; circummarginal grooves limited to distal ends of digits; supraarticular cutaneous folds slight; plantar surfaces of pes smooth, with prominently protuberant subarticular tubercles but lacking supernumerary tubercles; subarticular tubercles numbering three under Toe IV, two under Toes III and V, and one each under Toes I and II; decreasing digit length IV, III, V, II, I; outer metatarsal tubercle prominent and subcircular, pointed; inner metatarsal tubercle prominent and elongate, with a sharp venterolateral edge; digits of pes completely free of web; cloacal region smooth and devoid of supracloacaal tubercles or dermal flaps. Measurements of holotype: SVL 40.6; ED 5.6; TD 3.2; HL 17.3; SNL 7.7; IOD 3.9; HW 15.1; FA 8.6; TBL 24.5; TSL 13.5; PL 22.5; ML 10.3; Toe 4 L 15.0; Fin 1 L 5.9; Fin 3 L 7.2; Fin 3 DW 1.5; Fin 3 PPW 0.8; Toe 4 DW= 1.5; Toe 4 PPW= 0.6. Color of holotype in preservative: Dorsum homogeneous dark brown, blending laterally into light and dark brown marbled flanks with scattered enlarged distinct white spots; dorsal surfaces of head grayish; lateral surfaces of snout and head darker, nearly black with light brown tympanum and distinct white spots on lips; postrictal region heterogeneous, with black and brown blotches and white-tipped postrectal tubercles; dorsal surfaces of limbs brown with darker transverse bars (two on forearm, three on humerus, three on tibia, two on tarsus); dorsal surfaces of hands, feet, and digits dark brown with white bars on phalangeal articulations; inner two digits of hands and feet distinctly lighter than outer digital surfaces; venter pale cream; underside of throat speckled with brown, coalescing into nearly solid brown ventrolateral margins of throat; ventral surfaces of hands dark brown with prominent white subarticular tubercles; ventral surfaces of feet solid dark gray with light gray, velvety digital disks. Holotype color in life unrecorded. Var i at io n: There are three basic color patterns evident in our sample of P. adiastola from the type locality. The majority (9 of 11 specimens) exhibit solid gray dorsal coloration, with faint brown interoribital bars and only scattered flecks of black associated with dorsal tubercular ridges. This largely pigment-free pattern continues on to dorsal surfaces of limbs in a few specimens (SAMA R61907, 61909) but all remaining specimens have distinct transverse brown bars on limbs. The other two major color patterns include a specimen with a single, thick, vertebral yellow stripe (SAMA R 61908) and one with two dorsolateral pale lines (SAMA R 60259). Lateral surfaces of head with dark bar from tip of snout, through loreal region, encircling orbit, and following supratympanic fold posteroventrally to rictus of jaw. In most specimens (8 / 11), this dark coloration covers the majority of the lateral portions of the head. In three specimens (SAMA R61907, 61909, 60259), lateral portions of the head are distinctly lighter and darker pigmentation is limited to the canthus rostralis and supratympanic fold. Flank coloration varies from a simple dark (dorsally) to light (ventrally) fade (SAMA R60257, 60259) to a more distinct stratification between dark (above) and pale (below), with numerous moderately distinct pale round spots scattered across flanks. Ventral surfaces of the body are uniform pale cream with throat similarly colored or with a diffuse dark gray ventral surface of the entire head and throat (SAMA R61906, 61908, UPNG 9974 - 5). In these specimens, the chin and lower lips are distinctly lighter than surrounding throat. Palmar surfaces of the hands vary from very dark gray with white subarticular tubercles (e.g., UPNG 9974) to pale cream throughout their ventral surfaces (SAMA R 60257). Plantar surfaces of the feet are uniform light gray (SAMA R 60258) to dark gray with lighter subarticular tubercles (SAMA R 61906, R 61908, UPNG 9974). Color in life: Based on color images of paratype SAMA R 61907 before preservation. Dorsal surfaces are khaki to dark tan or brown (e.g. Fig. 5). The dark canthal and supratympanic bars are dark brown, tympanum is light brown, the loreal region is dark brown with gold flecks and the labial region grayish pink with gold bars. The iris is gold above and dark below below pupil and postrictal tubercles are tan with white tips. Bars on limbs are varying shades of brown on a lighter tan or brown background. The dorsal surfaces of the digits are brown with white on spots on digital articulations and the inner two digits on hands and feet and bright white. The throat is white and the venter is white anteriorly, fading to cream or yellow ventrally by groin. Ventral surfaces of limbs are yellow or pink. Size dimorphism: The single available female specimen of P. adiastola has a snoutvent length of 71.7 mm which is considerably larger than all of the nine available P. schmidti specimens examined by us (Talasea population: SVL 41.1���57.6; Table 1). This suggests to us the possibility that P. adiastola may have a greater extent of sexual size dimorphism than P. schmidti (P. a d i a s t o l a male/female SVL = 0.57; P. schmidti = 0.69) but larger sample sizes will be required to address this question in the future. Ecology and natural history: Platymantis adiastola was observed, collected, and recorded at Wanui Camp in primary forest (two hours walking distance from Sampun Village; 0 5 o 21.638 'S, 152 o 05.266'E), 300���500 m above sea level. This site was sampled for 14 days (14���28 March 2000). At Wanui the new species was subjectively characterized as 8���10 times more abundant than P. schmidti. In contrast, P. schmidti is clearly the most common form at lower elevations. Vocalizations confirming the sole presence of P. schmidti have been recorded or noted at numerous coastal localities, e.g. Kokopo and the vicinity of Rabaul, (Gazelle Peninsula), Sulu and Silali (foothills of the Nakanai Mountains) and, most significantly, at Talasea, the type locality of P. schmidti (SJR and JF pers. obs.). It was also abundant near Sampun Village (~ 50 m above sea level) and no P. adiastola were confirmed from this lowland site. Where the two species have been observed together, we have noted slight but clear microhabitat differences between the two forms (e.g. Foufopoulos & Richards in press): P. adiastola has been consistently observed calling from elevated perches (leaves and branches) as high as 2 m above the ground. In contrast, P. schmidti called predominantly from the ground, from low, herbaceous vegetation, or from stumps or fallen logs, consistently less than 1 m above the ground (Menzies 1982 a, SJR and JF pers. obs.). Distribution: Confirmed only at two localities (Fig. 1): near Sulu and Silali Villages, foothills of the Nakanai Mountains, West New Britain (Foufopoulos & Brown 2004) and Wanui, East New Britain (Foufopoulos & Richards in press). Advertisement calls: The advertisement call of P. adiastola is markedly distinct from that of P. s c h m i d t i in both temporal and spectral characteristics. It is, however, similar in general call structure (a rapidly amplitude-modulated train of notes) and so its impression on the human ear is similar. Because the call of the new species is one of the primary characteristics distinguishing the new species, we describe its vocalizations and compare them to those of P. s c h m i d t i in some detail, below. The following descriptions are based on 4 vouchered recordings of P. adiastola (2 from the Nakanai Mountains; 2 from Wanui) and 11 of P. s c h m i d t i (9 from the Nakanai Mountains; 2 from Wanui). The call of P. adiastola is a series of shrill, high-frequency tonal notes, delivered in a train of variable total duration and note repetition rate. One very distinctive characteristic of the call is the progressive shortening of inter-note intervals throughout the call, such that calls start slowly and gradually shift towards a more rapid note delivery rate towards the end of the note train. The effect on the human ear is a sound akin to that of a marble being dropped and bouncing, faster and faster, until it comes to rest after a final burst of rapid strikes (Fig. 7 A). In contrast, the calls of P. s c h m i d t i consist of a much more rapid, evenly-spaced train of dull, lower frequency, stridulating, vibrational notes, like the sound produced by slowly running a finger over the teeth of a comb or running a stick down a length of wood fence (Fig. 7 B). Menzies stated that the call of P. s c h m i d t i ���lacks musical quality and merely sounds like a stick being rubbed against wooden railings��� (Menzies 1982 a: 2). Individual male mean note repetition rate ([total number of notes ��� 1]/time from beginning of first note to beginning of last) was 4.0��� 4.5 (= 4.3 �� 0.4 SD; n = 4) notes/s for P. adiastola and 9.7���15.3 (= 12.8 �� 3.7 SD; n = 9) notes/s for P. s c h m i d t i. Calls of individual males of P. adiastola contained as few as 9���15 (mean 11.6 �� 3.6 SD) to as many as 11���38 (17.1 �� 8.9 SD) notes per call. The average number of notes per call for our sample of four males was 16.6 �� 3.4 (11.6���17.1; n = 5). Calls of individual male P. s c h m i d t i contained as few as 8���15 (mean 13.6 �� 5.2 SD) to as many as 42���55 (45.2 �� 8.1 SD) notes per call. The average number of notes per call for our sample of 13 P. schmidti recordings was 23.9 �� 8.8 SD (13.6���45.2). Individual note structure between the two species was quite different. The shrill, high frequency tonal notes of the call of P. a d i a s t o l a have a relatively complex internal temporal and spectral structure (Fig 8). Each note contains three distinct frequency components (Fig. 8 A, C) and a complex, amplitude modulated internal note structure with pronounced differences in call energy contained in the different amplitude modulated within-note pulses (Fig. 8 B). The dull, lower frequency twin-pulse notes of the call of P. schmidti (Fig. 9) are quite different in that each note contains a tightly-spaced twin-pulse structure (Fig. 9 B���D), with pulses only slightly differentiated in amplitude (Fig. 9 B, D) and both very similar to one another in spectral structure (Fig. 9 C). Menzies referred to the call of P. schmidti as "little more than a succession of double clicks��� (Menzies 1982 a: 2). In P. adiastola calls, each note contains a clear fundamental frequency component at 1.0 to 2.2 kHz (mean peak frequency = 1.1���1.3 kHz for four males), an emphasized (=dominant) frequency between 2.5 and 3.6 kHz (mean peak frequency = 3.0��� 3.2 kHz), and then an additional frequency component between 4.2 and 5.0 kHz (mean peak frequency 4.3���4.8 kHz). The three frequency components are stereotyped, with variation in frequency between males present (and apparently varying inversely proportionately with body size; correlations not shown) but with little or no within-male or within-call frequency modulation or energy shifts. In P. s c h m i d t i, the emphasized (= dominant) frequency lies between 2.2 and 2.9 kHz (peak frequency = 2.4���2.6 kHz for 11 males), and then minor frequency components include a fundamental between 1.1���1.9 kHz (= 1.5 kHz) and weak harmonics between 7.4 and 8.6 kHz. Unlike the new species, P. schmidti males exhibit a shift in emphasized frequency across the duration of long calls. Thus, calls typically start with the first 6���10 notes at a lower dominant frequency (2.4���2.6 kHz) and males shift the calling energy of subsequent notes to higher dominant frequencies (2.8���2.9 kHz) towards the end of the call. Individual notes of P. adiastola begin with 2���4 internal amplitude modulated pulses of 5���10 ms (Fig. 8 B), followed by the intense tonal note (Fig. 8 B���D) of 18���25 ms. Average relative intensity (in dB) of each of the preceding pulses is approximately 45���55 % of the subsequent tonal portion of each note. Total note duration for individual males ranged from 42.2���46.5 ms (= 44.8 �� 3.8; n= 13 notes) to 47.4���53.5 ms (= 50.8 �� 3.1; n= 17 notes); mean note duration for our entire sample of recordings was 47.3 �� 4.4 (42.3���56.7; n= 4). Inter-note intervals for the same males ranged from 232���255 ms to 245���267 ms; mean inter-note interval for the entire sample was 252.2 �� 26.3 SD ms (242.3���254.7; n= 4). Individual notes of P. schmidti are paired pulses (Fig. 8 B���D) of similar amplitude (Fig. 9 D). Both pulses begin with a rapid rise to peak amplitude (2���5 ms), followed by a more gradual decline to ambient levels (Fig. 9 B); in the second pulse, time for amplitude to fall to ambient levels is 2���3 times that of the first pulse. The average relative intensity (in dB) of the first pulses is approximately, Published as part of Brown, Rafe M., Richards, Stephen J., Sukumaran, Jeet & Foufopoulos, Johannes, 2006, A new morphologically cryptic species of forest frog (genus Platymantis) from New Britain Island, Bismarck Archipelago, pp. 45-68 in Zootaxa 1334 on pages 50-63, DOI: 10.5281/zenodo.174282, {"references":["Menzies, J. I. (1982 a) The voices of some male Platymantis species of the New Guinea region. British Journal of Herpetology, 6, 1 - 5.","Foufopoulos, J. & Brown, R. M. (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."]}