Sumaterana crassiovis, Arifin, Umilaela, Smart, Utpal, Hertwig, Stefan T., Smith, Eric N., Iskandar, Djoko T. & Alexander Haas,, 2018

Arifin, Umilaela, Smart, Utpal, Hertwig, Stefan T., Smith, Eric N., Iskandar, Djoko T. & Alexander Haas,, 2018, Molecular phylogenetic analysis of a taxonomically unstable ranid from Sumatra, Indonesia, reveals a new genus with gastromyzophorous tadpoles and two new species, Zoosystematics and Evolution 94 (1), pp. 163-193 : 169-174

Sumaterana crassiovis comb. n. Figs 2 Clade A, 5a, 6a

Rana pantherina Van Kampen, 1910.

Rana crassiovis Boulenger, 1920.

Rana (Hylorana) kampeni Boulenger, 1920.

Rana (Hylorana) crassiovis Boulenger, 1920.

Rana (Hylarana) kampeni Van Kampen, 1923.

Rana (Hylarana) crassiovis Van Kampen, 1923.

Rana (Chalcorana) kampeni Dubois, 1992.

Rana (Chalcorana) crassiovis Dubois, 1992.

Hydrophylax kampeni Frost et al., 2006.

Hydrophylax crassiovis Frost et al., 2006.

Hylarana kampeni Che et al., 2007.

Hylarana crassiovis Che et al., 2007.

Chalcorana kampeni Fei et al., 2010; Oliver et al., 2015.

Chalcorana crassiovis Fei et al., 2010; Oliver et al., 2015.

Syntypes.

Two adult females (BMNH1947.2.3.99 and BMNH1947.2.4.1-Fig. 7), Kerinci, Sumatra, Indonesia, 4000 feet (~1219 m a.s.l.), coll. Robinson-Kloss Expedition on the Batrachians. Based on the lack of morphological distinguishing characters ( Inger and Iskandar 2005) and low genetic divergence (2.56%, Suppl. materials 3) of topotypic specimens (this study), we consider C. kampeni a junior synonym of S. crassiovis comb. n..

Referred specimens (283).

262 adults (128 of them: 96 males and 32 females; were measured) and 21 tadpoles collected from Aceh up to Lampung (Appendix 1).

Description.

Specimens were assigned to Sumaterana crassiovis comb. n. based on comparison of material from Kabupaten Kerinci. Sumaterana crassiovis comb. n. is described by the following combination of characters: a medium sized species, SVL in males 30.03-48.87 mm, females 40.98-83.99 mm; head width subequal to head length; snout rounded, obtusely pointed in dorsal view, slightly protruding in lateral view; nostril closer to snout than to eye; vomerine teeth present, in oblique groups, between choanae; tongue lanceolate; loreal area deeply concave; canthus rostralis sharp, constricted behind nostrils; rictal ridge present; tympanum distinct, translucent (not transparent); interorbital distance 75.96-124.80% width of upper eyelid in females, 68.26-120.31% width of upper eyelid in males; pineal spot visible; dorsolateral fold absent; supratympanic fold thick, posttympanic fold absent; dorsum finely granulated with scattered tubercles, variable in size and density; flanks coarsely granulated with few tubercles; venter smooth, granulated posteriorly; rear of thigh usually barred as continuation of thigh dorsal pattern; arm slender, lower arm length 19.03-24.18% SVL in females and 19.58-25.46% SVL in males; hand length 31.54-36.98% SVL in females and 31.77-39.23% SVL in males; fingers long, without webbing; fingertips expanded into discs, diamond-shaped, with circummarginal groove; Finger I <Finger II, Finger III longest; fringes present on the outer phalanges of all fingers; subarticular tubercles distinct; width of Finger III disc> width of Toe IV disc; hindlimbs long, articulation of the heels reaching beyond tip of snout, when limb aligned to body; relative femur length 85.39-94.32% tibia length in females, 85.82-95.02% tibia length in males; length of tibia 60.17-70.52% SVL in females, 58.78-76.44% in males; toes slender and long; tip of toe extended into disc, diamond-shaped, with circummarginal grooves; toe lengths: I <II <III <V <IV, Toe V only slightly longer than Toe III; Toes I, II, III, and V fully webbed, webbing of Toe IV usually one phalanx free (I(1+/-―1+/-)II(1+/-―1+/-)III(1+/-―2+/-)IV(2+/-―1+/-)V); subarticular tubercles distinct; inner metatarsal tubercle distinct, oval, 92.07-212.77% T4DW in males and 98.80-150.00% in females; outer metatarsal tubercle absent; tarsal fold absent. (Measurements: Tables 4-5).

Coloration.

Dorsal skin background green in life, with dark blotches around tubercles, lighter areas on the dorsum forming irregular network pattern; dark line connects the eye and the snout; the upper and lower lips with dark blotches on a light background; iris golden yellow, reddish anteriorly and posteriorly, with a dark netting pattern; tympanum pale brown, encircled by a dark line; flanks lighter than dorsum, lighter ventrad and with dark spots; venter whitish, throat and chest with or without dark marking; distinct cross-bars on dorsal limbs; the rear of thigh with dark vertical bars (usually a continuation from dorsal surface and separated by narrow lighter areas) or mottling (dark marking on lighter background); ventral legs are dusted with brown pigment; webbing color brown. In preservative, dorsal background light brown; flanks becoming gray; iris changed to gray.

Variation.

(1) number of tubercles on dorsum and flanks: few to dense; (2) size of tubercles on dorsum: small and round to larger and elongated; (3) dorsolateral fold absent, but row of few small tubercles form incomplete dorsolateral series, dorsal to the posterior of trunk (not in continuation of tympanic fold); (4) dorsal coloration: dark blotches on green background vary from few and isolated, to dense, and forming irregular green background network between the dark blotches; (5) flank color yellowish-green to green (as dorsum), lighter ventrad, with distinct spots; (6) upper and lower lips: whitish to greenish, with dark markings, small distinct bars to wide and connected, lip markings absent or very thin in few individuals; (7) ventral dark markings: from none (ventral side whitish) to dark on throat and reaching venter, pale to dark; (8) rear of thigh with dark bars, complete or broken, or occasionally dark mottling on whitish/grayish background (Fig. 4 a–d); (9) iris: golden to pale yellow, from faint and thin to dense and dark netting; (10) number of cross bars: 3-4 on lower arm (from elbow to wrist), 4-7 on thigh; (11) Toe IV: from one phalanx free of webbing to webbing reaching intercalary tubercle of Toe IV (Fig. 4 e–h). See Fig. 8 for images of Sumaterana crassiovis comb. n. from different localities and for morphometric variation Tables 4-5.

Sexual dimorphism.

Males significantly smaller than females. Tympanum diameter 45.27-71.68% ED in males, 33.33-48.51% ED in females. Male with distinct undi vided nuptial pads, covering base of the first finger to subarticular tubercle in dorsal and medial surface, paired subgular vocal sacs, humeral glands absent.

Common name.

We propose Kerinci Cascade Frogs as the common English name (to replace the old spelling in "Korinchi Frog", Iskandar and Mumpuni 2004) and Katak Jeram Kerinci as the Indonesian name.

Distribution and ecological remarks.

This species is widespread on the island of Sumatra, ranging from the northern part of Provinsi Aceh to Kabupaten Pasawaran, the southern part of Provinsi Lampung (Fig. 9). Elevational range 425-1545 m a.s.l.. This species is abundant along rocky streams (usually 1-5 m wide) in primary or good secondary forest. The inhabited streams are typically rocky with boulders (usually diameter> 1 m) and with rock formations along the stream, water current velocity 0.2-1.1 m/s (Fig. 10). Males of this species commonly can be observed perching on rocks or vegetation at the stream banks. Females were rarely observed near the streams. It seems that they approach the streams only during breeding activities. Tadpoles were often found in groups, on rocks in the stream, overflown with water in cascading sections.

Tadpoles.

Tadpoles were identified (100%) using 12S rRNA+16S rRNA+tRNAval barcoding with adult samples from the type locality. We examined total of 21 tadpoles. Stage 25: MZB.AMPH.29362 (n = 1), ZMH.A12649 (n = 3), MZB.AMPH.29363 (n = 1), MZB.AMPH.29364 (n = 1), Stage 26: MZB.AMPH.29362 (n = 2), MZB.AMPH.29356 (n = 1), ZMH.A12649 (n = 2), MZB.AMPH.29365 (n = 1), Stage 31: MZB.AMPH.29362 (n = 1), Stage 32: MZB.AMPH.29362 (n = 1), Stage 35: MZB.AMPH.29362 (n = 1), ZMH.A12650 (n = 1), Stage 36: MZB.AMPH.29355 (n = 1), ZMH.A12649 (n = 1), Stage 39: MZB.AMPH.29360 (n = 1), Stage 42: MZB.AMPH.29361 (n = 1). One selected tadpole from the lot had 100% match (12S rRNA+16S rRNA+tRNAval) to an adult Sumaterana crassiovis comb. n. from the type locality. We refer to ZMH.A12650 (stage 35, Fig. 11 a–c) for tadpole description.

Head and trunk approximately oval in dorsal view and dorsoventrally depressed and streamlined, in lateral view; maximum body width 64.40% body length; snout expanded and broadly rounded with emargination laterally setting off snout from body; eyes positioned dorsolaterally, oriented laterally; ED = 2.31 mm; IND/IOD = 48.22%; SN/EN = 44.82%; nostril open without raised rim; positioned anterodorsally and anterolaterally directed; two glands clusters present, infraorbital glands (five on each side) and postorbital glands (one on each side); oral disc ventral, a groove separating upper lip from snout, ODW/BW = 66.33%; oral disc marginal papillae short, arranged in single row; marginal papillae of upper lip present only on sides, on lower lip in uninterrupted row; two short submarginal papillae in lateral area of upper lip; LTRF: 9(6-9)/9(1); upper jaw sheath broad and heavily keratinized, smooth, undivided, thick but with distinct thinner medial section; lower jaw sheath undivided, V-shaped, smooth, and thick; both jaw sheaths finely serrated along their edges; very large abdominal sucker adjoining oral disc posteriorly, SUL/BL = 76.61%, SUW/BW = 89.03%; spiracle sinistral, tube long and posterior half free from body wall, opening directed posteriorly or posterodorsally; anal tube median, free from tail fin, directed posteriorly; strongly muscular tail: TAL/BL = 165.71%, TMH/BH = 71.87%, TMH/MTH = 63.00%; upper fin convex; maximum upper fin height is 30.57% maximum tail height at 49.19% of tail length; tail tip pointed.

In life (Fig. 11 a–c), dorsum light brown, orangeish anteriorly and posteriorly to eyes; trunk darker than head; tail muscle light brown with fine-orange stippling; lower flanks region whitish; lateral tail vein very obvious, including dorsal branching along myosepta; upper and lower fins mostly transparent without iridophores; iris black, with dense gold to orange iridescent stippling; abdomen whitish laterally and densely stippled with fine-orange iridophores medially; abdominal sucker mostly transparent with white iridocytes in the center. In preservation, upper side gray with dark stippling; dense-dark stippling laterally; iris black; lens gray; ventral side uniformly transparent with some grey pigments in the anterior region of snout and lateral parts.