Limnonectes nephophilus Dehling, Neokleous, Das, Grafe, Min & Hertwig, 2025, sp. nov.

Limnonectes nephophilus sp. nov. Rana palavanensis (nec Rana palavanensis Boulenger, 1894): Inger 1985: 52 (partim). Holotype. NMBE 1056394, adult male, from Camp 3 (4.038079°, 114.888078°; 1,350 m), Gunung Mulu National Park, Sarawak, Malaysia, collected in March 2009 by J. M. Dehling, N. W. Tinggang, Y. M. Pui, S. T. Hertwig and A. Jankowski (Fig. 3D). Paratypes. BMNH 1978.214 – 215, two adult males, collected on 7–8 September 1977 at the type locality by J. Dring; BMNH 1978.1666 – 1667, two adult females, BMNH 1978.1668 – 1669, two juveniles, all from the type locality, collected in 1978 by J. Dring. NMBE 1065176–1065177, two tadpoles, from Camp 4 (4.044178°, 114.913847°; 1,800 m), Gunung Mulu National Park, Sarawak, Malaysia, collected on 29 March 2007 by S. T. Hertwig and K. Lilje. NMBE 1061685, adult female, NMBE 1061686, adult male, collected on 24 February 2013 by S. T. Hertwig, A. Jankowski, Y. M. Pui; NMBE 1073768, adult male, NMBE 1073755, adult female, collected on 6 March 2016 by J. Flury, A. Haas, S. T. Hertwig, Y. M. Pui and N. Reichen; all from Lepo Bunga (3.941808°, 115.544667°; 1,760 m), Gunung Murud, Pulong Tau National Park, Sarawak, Malaysia. NMBE 1061641–1061642, two juveniles, collected on 29 February 2012 by S. T. Hertwig, A. Jankowski and Y. M. Pui; NMBE 1073581, adult male, NMBE 1073590, adult female, NMBE 1073706, 1073709, two juveniles, collected on 9–12 March 2016 by J. Flury, A. Haas, S. T. Hertwig, Y. M. Pui and N. Reichen; all from Church Camp (3.926317°, 115.513889°; 2,100 m), Gunung Murud, Pulong Tau National Park, Sarawak, Malaysia. NMBE 1073578, adult male, from Summit Trail (2,100 m), Church Camp, Gunung Murud, Pulong Tau National Park, Sarawak, Malaysia, collected on 10 March 2016 by J. Flury, A. Haas, S. T. Hertwig, Y. M. Pui and N. Reichen. NMBE 1073715, unsexed adult, from Lepo Bunga (3.941808°, 115.544667°; 1,760 m), Gunung Murud, Pulong Tau National Park, Sarawak, Malaysia, collected on 7 March 2016 by J. Flury, A. Haas, S. T. Hertwig, Y. M. Pui and N. Reichen. Referred material. BMNH 1978.216, tadpole, from the type locality, collected on 8 September 1977 by Julian Dring. Diagnosis. Limnonectes nephophilus sp. nov. belongs to the L. palavanensis species group, showing the typical characters of the group, including small size (SVL <40 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum finely shagreened, with prominent longitudinal dorsolateral glandular ridges; interorbital distance equal to or larger than upper eyelid width; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from all other species of the group by a unique combination of morphological characters, including toe webbing very reduced; supratympanic fold subangular with anterior part shorter than posterior part; pineal spot distinctly discernible, slightly anterior to level of anterior upper-eyelid edge; vomer ridges short, widely separated from each other; median ridge low, indistinct, running from level of nostrils to level of pelvis; postpalpebral and lumbar ridges absent; size relatively large with SVL of adult males 26.1–31.9 mm, of adult females 33.1–37.6 mm. The species differs from all other species of the group by at least 5.7% in the sequence of the 16S barcoding gene. The advertisement call is a sharp chirp, 75–90 ms in duration at a dominant frequency of 2.5–3.0 kHz with a marked frequency modulation, given in call bouts of up to 21 calls with an intercall interval of 135–383 ms. The species occurs in montane rainforest between 1,200 and 2,100 m. Description of holotype. Adult male; SVL 28.4 mm; body moderately sturdy, widest at temporal region, slightly tapering to groin (Fig. 3D); head large (HL/SVL 0.37, HW/SVL 0.38), about as wide as long (HW/HL 1.03); snout moderately long (SL/HL 0.40), rounded in both dorsal view and profile, slightly projecting beyond lower jaw, wider than long (SL/EE 0.82); canthus rostralis distinct between eye and nostril, slightly concave; loreal region oblique, moderately concave; nostrils rounded, directed laterally, situated slightly closer to tip of snout than to eye (EN/NS 1.09), separated from each other by distance much larger than distance between eye and nostril (NN/EN 1.70); eye directed anterolaterally, moderately protruding, large (ED/HL 0.32), its diameter shorter than snout (ED/SL 0.80); interorbital distance greater than upper eyelid width (IO/EW 1.19) and smaller than internarial distance (IO/NN 0.69); pineal spot distinctly discernible, slightly anterior to level of anterior upper-eyelid edge; tympanum and its annulus distinctly visible, except posterior one-fourth which is covered by thick skin; tympanum separated from eye by little more than one-third of its diameter (ET/TD 0.37); tympanum diameter smaller than eye diameter (TD/ ED 0.61); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, rounded, located far anterolaterally at margins of roof of mouth, covered by palatal shelf of maxilla in ventral view; vomer processes bearing teeth, short, separated from each other and from choana by length of individual process (Fig. 4D); tongue moderately long and narrow, bilobed for about one-sixth of its length, free distally for about half its length; median lingual process absent; vocal sac apertures slit-like, directed posterolaterally, situated closer to corner of mouth than to base of tongue. Dorsal surfaces of head, trunk and limbs finely shagreened; lateral surfaces of trunk finely areolate with scattered small tubercles; dorsolateral ridge prominent, from posterior end of upper eyelid to groin; median ridge low, indistinct, running from level of nostrils to level of pelvis; postpalpebral and lumbar ridges absent; horseshoe-shaped dermal ridge low, in middle of back just behind level of arms; supratympanic fold thick and conspicuous, extending from posterior end of orbit to insertion of arm, subangular with anterior part shorter than posteroventrally directed posterior part (Fig. 5D); row of small tubercles on lateral edge of upper eyelid, postaxial side of forearm, and lateral part of head; ventral side of head smooth with weak dermal folds parallel to posterior part of mandible; weak transverse chest fold between arm insertions; ventral side of trunk weakly areolate with sparsely scattered small tubercles; ventral side of limbs smooth; cloaca without dermal flap. Forelimbs moderately sturdy; hand relatively small (HND/SVL 0.24); tips of fingers rounded, slightly enlarged into disks; relative length of fingers: II = IV <I <III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV larger and more prominent than distal ones; finger webbing absent; thenar tubercle distinct, oval, flat, little more than one-third length of metacarpal of Finger I; inner palmar tubercle indistinct, small, rounded, flat, on proximal end of metacarpus of Fingers II and III; outer palmar tubercle indistinct, flat, elongated, on proximal half of metacarpal of Finger IV; nuptial pads absent. Hindlimbs sturdy, very long (LEG /SVL 1.84); heel reaching to tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.57), slightly longer than thigh (TFL/THL 1.11); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot slightly shorter than tibiofibula (FOT/TFL 0.93); relative length of toes: I <II <V <III <IV; toe tips rounded, slightly enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1.5/2.25 II 1.5/3- III 2.25/3.25 IV 3.25/2+ V (Fig. 6); dermal flap on postaxial side of Toe V, widest from proximal end of metacarpus to proximal subarticular tubercle, continued to disk; inner metatarsal tubercle very prominent, elongated, half as long as metatarsus of Toe I; outer metatarsal tubercle absent; low ridges of callous tissue from proximal plantar part of metatarsus to proximal subarticular tubercle on Toes I, II, and V. Colouration. In life (Fig. 7D), dorsum of holotype including lateral parts light brown with orange tinge; interorbital bar blackish brown; head anterior to interorbital bar as well as tympanic area bright orange; horseshoe-shaped tubercle slightly darker brown than dorsum and dark brown in the centre; two large black dots in longitudinal row at level of pelvis; area around cloaca dark brown; supratympanic fold dark grey; dorsolateral fold red in anterior part, lightening to orange posteriorly, bordered laterally by thin black line; tympanum light brown; side of snout light brown with larger dark brown spots and dark bar along canthus rostralis; arms orange, darkening to light brown towards hands; lower arm with dark brown crossbands; hands yellowish brown; legs reddish light brown with greyish light brown crossbands; tarsus and toes red; ventral side of head, chest, arms, lower legs and tarsus yellowish bright orange; large dark brown spots in infralabial region; throat speckled with minute brown dots; abdomen bright yellow, darkening laterally, with white speckling; thighs dark orange, darkening to reddish preaxially and postaxially; preaxial side of thighs with bluish white speckling; plantar side of feet blackish. In preserved state (Fig. 3D), dorsum creamish light brown; arms, fingers and toes as well as dorsal part of snout yellowish light brown; upper eyelids light blue; legs reddish light brown; upper arms without crossbands; lower arms, fingers, legs including feet with regular dark brown crossbands; interorbital bar blackish brown; supratympanic fold greyish brown; dorsolateral fold bordered laterally by thin black line; tympanum light brown; three large dark brown spots below eye and nostril on each side; horseshoe-shaped tubercle slightly