Rana morafkai, BAIN & LATHROP & MURPHY & ORLOV & CUC, 2003

BAIN, RAOUL H., LATHROP, AMY, MURPHY, ROBERT W., ORLOV, NIKOLAI L. & CUC, HO THU, 2003, Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species, American Museum Novitates 3417, pp. 1-60 : 44-46

Rana morafkai , new species

(Previously referred to as species 5, ‘‘Mottled’’)

Figures 12K, L , 13E , 14 B–D

HOLOTYPE: (ROM field no. 7301) ROM 39932, a female from Tram Lap , An Khe District, Gia Lai Province, Vietnam (14°26̍39̎N, 108°32̍97̎E, elevation ca. 900 m) collected on 19 June 1996 by R.H. Bain, B. Hubley, A. Lathrop, R.W. Murphy, and N.L. Orlov. The holotype had leg and liver tissue removed shortly after it was euthanised.

PARATYPES: ROM 39904–39911, 39930, 39934, 39937, 39947, and 39949 collected with the holotype between 15 and 19 June 1996 by R.H. Bain, B. Hubley, A. Lathrop, R.W. Murphy, and N.L. Orlov; ROM 39904– 39011, and 39937 are males; the remainder are females . ROM 25094–25097, 25099, 25101, 25104 –25106, 25108–25111, all males collected in Buon Loi , An Khe District, Gia Lai Province, Vietnam (elevation 700–750 m) by I.S. Darevsky and N.L. Orlov on 11 November 1993 .

DIAGNOSIS: Rana morafkai , a member of the subgenus Odorrana (sensu Fei et al., 1990), is characterized by a combination of the following attributes: (1) body dorsoventrally compressed; (2) SVL means of males 43 mm (39–46 mm), females 88 (80–100 mm); (3) vomerine teeth in rows oblique to choanae; (4) yellow­white lip­stripe extending across upper lip, terminating in glandule above insertion of arm; (5) head broad, bluntly rounded in profile; (6) tympanum round, relatively large, distinct, approximately 96% of eye length in males, 71% in females; (7) supratympanic fold weak; (8) dorsal skin smooth or partly shagreened, becoming granular laterally, dorsolateral folds absent; (9) dorsum changing colors between day and night, usually bright green in daylight, brown at night sometimes with black spots; forelimbs and hindlimbs with transverse bars; (10) median callous pad on finger III to proximal tubercle; (11) disks on fingers and toes greatly enlarged (>2× base of phalanges); (12) webbing on feet complete to base of disk in females, sometimes as a fringe on IV; in males to distal tubercle in males, lateral fringes on I and V to terminal phalanges; webbing brown­gray; (13) subarticular tubercles and inner metatarsal tubercle conical, indistinct; (14) terminal phalanges T­ shaped; (15) xiphisternum deeply notched posteriorly; (16) males with nuptial pads on thumb, paired gular pouches, pectoral spines absent; (17) eggs white.

COMPARISONS: Rana morafkai resembles other Asian cascade ranids, including Huia nasica , Rana andersonii , R. archotaphus , R. bacboensis , R. chalconota , R. chloronota , R. daorum , R. grahami , R. graminea , R. hainanensis, R. hejiangensis , R. hmongorum , R. hosii , R. jingdongensis , R. junlianensis , R. kwangwuensis , R. leporipes , R. livida , R. margaretae , R. schmackeri , R. sinica , and R. tiannensis (table 12). It differs from these cascade ranids (except R. chalconota ) in having unique nocturnal and diurnal coloration, less distinct subarticular tubercles (all other species bear distinct subarticular tubercles), and a webbing pattern that is unique to each sex (females fully webbed to disk, males to distal subarticular tubercles). Female R. morafkai (SVL 80–100 mm) are considerably larger than female R. archotaphus (59–62 mm), R. chalconota (45–60 mm), R. daorum (55–58 mm), and R. sinica (holotype 66.6 mm). The presence of a large tympanum in the males (TMP:EYE 0.96) differentiates R. morafkai from H. nasica (0.5), R. andersonii (0.7), R. chloronota (0.57), R daorum (0.29), R. grahami (0.53), R. graminea (0.77), R. hainanensis (0.66), R. hmongorum (0.43), and R. hejiangensis (0.5). Its broadly rounded snout distinguishes R. morafkai from H. nasica , R. andersonii , R. chalconota , R. schmackeri (obtusely pointed snouts in profile), R. graminea , and R. margaretae (distinctly depressed snouts). The presence of gular pouches differentiates male R. morafkai from those of R. andersonii , R. chalconota , R. grahami , R. hainanensis , R. hmongorum , R. jingdongensis , R. hosii , and R. margaretae . The absence of dorsolateral folds distinguishes R. morafkai from R. chalconota , R. daorum , R. graminea , R. hosii , R. junlianensis , R. kwangwuensis , R. leporipes , and R. margaretae (pustules on the dorsum of R. grahami and R. hmongorum sometimes form a dorsolateral fold). The absence of an outer metatarsal tubercle distinguishes R. morafkai from R. archotaphus and R. chalconota . The presence of completely white eggs differentiates it from R. bacboensis (black eggs), H. nasica , R. andersonii , R. chalconota , R. grahami , R. junlianensis , R. margaretae , and R. schmackeri (white eggs with a dark melanic pole). The absence of spinules on the venter distinguishes R. morafkai from R. andersonii , R. grahami , R. jingdongensis , R. junlianensis , R. margaretae and R. schmackeri . Rana sinica differs from R. morafkai by its indistinct and skincovered tympanum (distinct and uncovered in R. morafkai ), finger formula (I <II <IV for R. sinica, II <I <IV for R. morafkai ), rounded distal phalanges (T­ shaped in R. morafkai ), and absence of a lip­stripe. Rana leporipes has small disks and white supratympanic folds, both distinguishing it from R. morafkai . The color pattern and skin texture of specimens of R. morafkai may be identical to those of R. chloronota ; however, the pronounced sexual dimorphism in the tympanum size (TMP:EYE R. morafkai males 0.96; females 0.71) differentiates it from all Vietnamese populations of Rana chloronota except those from Na Hang (males 0.60, females 0.43).

DESCRIPTION OF THE HOLOTYPE: An adult female (ROM 39932), head width 66% of length, length 50% of SVL; snout short, acutely rounded in dorsal view, bluntly rounded in profile, protruding beyond margin of lower jaw; eye large, prominent, 38% of snout length; eyelid broader than interorbital distance. Top of head flat; canthus rostralis rounded; loreal region concave; lip flared just anterior to orbit; nostril about three­fourths distance from eye to tip of snout; supratympanic fold curving posteroventrally from posterior corner of eye to a level above the insertion of the arm; tympanum round, distinctly visible, separated from eye by distance equal to TMP, 68% of EYE. Choanae ovoid; vomerine dentigerous processes prominent, oblique, posteromedial to choanae, each bearing numerous teeth. Tongue cordiform, distinctly notched posteriorly, free for approximately two­thirds its length.

Forearms moderately robust; fingers moderately short, slender, hands 21% of SVL, relative lengths of fingers II <I <IV <III, ventromedial callous pad on finger III to proximal tubercle, disks greatly expanded (>2× base of phalanges), relative pad size II <I <IV <III, pad width (III) 90% of pad length, ventral circummarginal grooves present; terminal phalanges T­ shaped; subarticular tubercles conical. Hindlimbs moderately robust; tibia length 62% of SVL; foot length 82% of SVL; relative toe lengths I <II <III <V <IV; inner tarsal fold absent; feet fully webbed to toe disk, lateral fringe on I and V to terminal phalanges, toes long, slender, with large, rounded, triangular disks, relative pad size I = II = III> IV k V, pad width (IV) 80% of pad length, each with ventral circummarginal grooves; subarticular tubercles prominent and conical; inner metatarsal tubercle ovoid, long; outer metatarsal tubercle absent.

Xiphisternum large, deeply notched posteriorly.

Skin on dorsum shagreened, light granulations laterally; dorsolateral folds absent; small tubercles posteroventrally to tympanum; prominent granules on flanks and around cloaca; cloacal opening unmodified, directed posteriorly at upper level of thighs.

COLOR IN LIFE (in preservative): Green (livid blue, with dark spots); flanks yellow and gray (cream and gray) with some white mottling posteroventrally; lip­stripe yellowwhite (creamy white); loreal region brown (black); tympanum beige with dark brown central ring; limbs above brown with black transverse bands, below creamy yellow with black mottling; thighs marbled yellow and brown; cloacal region black; venter creamy white; iris golden, margin of pupil outlined in a striking yellow and red border.

SECONDARY SEXUAL CHARACTERS: The holotype is a gravid female with creamy white eggs, 2 mm in diameter. Gravid females are approximately twice the SVL of males. Tympanum in males (TMP:EYE 0.96) is larger than in females (0.71). Webbing on females extends to disk, and only to distal subarticular tubercle in males. Males have velvety nuptial pads extending across the thumb, paired gular pouches located at the angle of the jaws, and pectoral spines absent.

MEASUREMENTS OF HOLOTYPE (in mm): SVL 84.4; SNT 14.3; HDL 42.7; HDW 28.2; EYE 5.4; IOD 7.3; TMP 3.7; TEY 4.1; HND 18.0; FGR 14.0; FPL 3.0; FPW 2.7; TIB 52.6; FTL 68.7; TPL 2.9; TPW 2.3.

VARIATION OF PARATYPES: The skin on the dorsum can be smooth or partly shagreened. These frogs are often brown at night, but diurnally they can become green over their entirely body or only dorsally. The dorsum is occasionally colored with black spots. Variation in all type material is given in table 15.

MEASUREMENTS OF FEMALE PARATYPES (in mm, n = 4, ROM 39930, 39934, 39947, 39949): SVL 87.6 ± 6.9 (80.4–99.6); SNT 13.4 ± 1.7 (10.8–15.6); HDL 39.1 ± 2.4 (37.6–38.5); HDW 27.2 ± 1.1 (27.7–28.2); EYE 6.0 ± 0.9 (4.9–7.6); IOD 7.7 ± 0.9 (6.1–9.0); TMP 4.1 ± 0.7 (3.4–5.2); TEY 4.5 ± 1.9 (3.2–8.3); HND 18.8 ± 2.2 (16.7– 21.9); FGR 15.0 ± 2.5 (12.2–18.1); FPL 3.5 ± 0.4 (2.8–3.9); FPW 3.0 ± 0.3 (2.7–3.5); TIB 53.5 ± 4.7 (47.4–61.5); FTL 63.0 ± 11.3 (43.3–76.1) TPL 3.1 ± 0.3 (2.5–3.5); TPW 2.9 ± 0.5 (2.3–3.7).

MEASUREMENTS OF MALE PARATYPES (in mm, n =14, ROM 25094–25097, 25099, 25101, 25104–25106, 25108–25111, 39937): SVL 43.2 ± 1.7 (39.2–45.9); SNT 6.8 ± 0.4 (6.2–7.8); HDL 21.3 ± 0.8 (20.5–22.4); HDW 13.8 ± 1.0 (13.2–15.5); EYE 3.5 ± 0.5 (2.5–4.5); IOD 3.9 ± 0.4 (3.2–4.8); TMP 3.3 ± 0.3 (2.8–4.0); TEY 1.2 ± 0.3 (0.8– 1.8); HND 11.8 ± 1.4 (9.0–15.6); FGR 9.6 ± 0.8 (7.9–12.0); FPL 1.8 ± 0.3 (1.16–2.3); FPW 1.6 ± 0.2 (1.4–2.0); TIB 26.0 ± 1.5 (22.8–28.4); FTL 27.1 ± 7.7 (17.5–40.9); TPL 1.7 ± 0.2 (1.3–2.0); TPW 1.3 ± 0.1 (1.1–1.7).

ETYMOLOGY: The specific name honors David Joseph Morafka, Research Associate of the Royal Ontario Museum and California Academy of Sciences, in recognition of his unfailing friendship, his unselfish development and perpetuation of multiple, independent research programs, and for his catapulting the careers of many conservation biologists.

DISTRIBUTION AND ECOLOGY: Rana morafkai is known only from the Tay Nguyen Plateau of the Central Highlands , Gia Lai Province, Vietnam. It inhabits forested montane river systems. Specimens may be found on or near rapids or waterfalls. In at least May and June, male specimens may have distended gular pouches, indicating that this species breeds in spring .

REMARKS: This species has been previously referred to as R. livida by Inger and Chanard (1997) and Inger et al. (1999).

Figures

Fig. 12. Heads of members of the Rana chloronota complex in lateral and dorsal views: (A, B) Rana chloronota, male (ROM 26402), and (C, D) female (ROM 26406); paratypes of (E, F) Rana bacboensis, new species (ROM 29529); (G, H) Rana daorum, new species (ROM 38538); (I, J) Rana hmongorum, new species (ROM 39891); (K, L) Rana morafkai, new species (ROM 39906); (M, N) Rana banaorum, new species (ROM39941); and (O, P) Rana megatympanum, new species (ROM 26398).

Fig. 13. Hands and feet of members of the Rana chloronota complex: (A) Rana chloronota (ROM 26402); paratypes of (B) Rana bacboensis, new species (ROM 29529); (C) Rana daorum, new species (ROM 38538); (D) Rana hmongorum, new species (ROM 26379); (E) Rana morafkai, new species (ROM 39906); (F) Rana banaorum, new species (ROM 39941); and (G) Rana megatympanum, new species (ROM 26398). Scale equals 5 mm.

Fig. 14. Members of the Rana chloronota complex from Vietnam. A. Paratype of R. hmongorum, new species, male from Lao Cai Province. (B) Female, brown (night) phase, (C) female, green (day) phase, and (D) male of R. morafkai, new species, from Gia Lai Province. (E) Female and (F) male of R. banaorum, new species, from Gia Lai Province. (G, H) female of R. megatympanum new species.