Rana daorum, 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 : 38-40

Rana daorum , new species

(Previously referred to as species 4, ‘‘Small’’)

Figures 11E, F , 12G, H , 13C

HOLOTYPE: (ROM field no. 19047) ROM 26381 an adult female from approximately 5 km NW of Sa Pa village , near O Qui Ho Pass, Lao Cai Province, Vietnam (22°22̍09̎N, 103°50̍14̎ E, elevation 1400 m) collected on 7 May 1995 along a waterfall beside the road by A. Lathrop and B. Hubley at approximately 1930 hours.

PARATYPES: ROM 26382–26397, all males, collected with the holotype, 7 May 1995 by A. Lathrop and B. Hubley at 1030 hours ; ROM 38500–38530, 38532–38540, 38542– 43, 38546–38561 collected between 30 April and 15 May 1999 in the vicinity of the type locality approximately 12 km northwest of Sa Pa Village near the O Qui Ho Pass (22°20̍58̎N, 103°46̍14̎E, elevation 1900 m) by R.O. de Sa´, C.T. Ho, A. Lathrop, T. Mason, R.W. Murphy, and N.L. Orlov . ROM 38547 is a subadult ; ROM 38503, 38507, 38512, 38516, and 38530 are gravid females ; ROM 38500, 38517, 38526, and 38538 are nongravid females; and the remaining specimens are males with distended gular pouches .

DIAGNOSIS: Rana daorum , 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 36 mm (32–38 mm), females 55 mm (53–58 mm); (3) vomerine teeth absent; (4) lip­stripe white, extending across upper lip, terminating in a glandule above insertion of arm; (5) head not broad, snout rounded in dorsal view, rounded in profile; (6) tympanum round, distinct, TMP:EYE of females (0.45) greater than males (0.29); (7) supratympanic fold weak or absent; (8) dorsal skin smooth, granular near cloaca and tympanum, dorsolateral fold covered with small white granules, ventral skin smooth; (9) dorsum green occasionally with black spots, flanks brown with at least one large white glandular spot; fore­ and hindlimbs goldish brown, with mottling or indistinct transverse bands; (10) median callous pad on proximal two­thirds of fingers II and III; (11) disks on fingers and toes greatly enlarged (>2× base of phalanx), finger pads larger than toe pads; (12) feet fully webbed to disks, but as a fringe from distal subarticular tubercle of IV, lateral fringes on I and V to terminal phalanges, webbing mottled brown; (13) subarticular tubercles and inner metatarsal tubercle distinct, conical; (14) terminal phalanges T­ shaped; (15) xiphisternum narrow, forked posteriorly; (16) males with nuptial pads on thumb, paired gular pouches, pectoral spines absent; (17) eggs large, white.

COMPARISONS: Though bearing a superficial resemblance to other cascade ranids of Southeast Asia, R. daorum is distinct (table 12) by the absence of vomerine teeth, females with larger TMP:EYE than males, and the presence of at least one large white spot on each flank. Its noticeably smaller SVL, dorsolateral folds formed by white granules, and solid, bright green dorsum immediately distinguishes it from H. nasica , R. andersonii , R. archotaphus , R. bacboensis , R. chloronota , R. grahami (whose dorsal pustules sometimes form a fold), R. graminea , R. hainanensis , R. hejiangensis , R. jingdongensis , R. junlianensis , R. kwangwuensis , R. livida , R. margaretae , R. schmackeri , R. sinica , and R. tiannensis . The entirely white eggs of R. daorum distinguish it from Huia nasica , Rana andersonii , R. chalconota , R. grahami , R. junlianensis , R. margaretae , R. schmackeri (all with white eggs with a melanic pole), and R. bacboensis (fully pigmented eggs). The presence of gular pouches in males differentiates R. daorum from R. andersonii , R. chalconota , R. grahami , R. hainanensis , R. hosii , R. jingdongensis , R. junlianensis , R. kwangwuensis , and R. margaretae . A distinct and uncovered tympanum also distinguishes R. daorum from R. sinica . Rana daorum can be further distinguished from R. livida , R. chloronota , R. maragaretae , and R. schmackeri by its finger formula (II <I <IV for R. daorum , I <II <IV for others listed). Although R. daorum closely resembles A. chunganenesis in size, the presence of white granular dorsolateral folds, and diurnal behavior, A. chunganensis is red­ brown, has vomerine teeth, and males have a TMP:EYE ratio twice that of R. daorum .

DESCRIPTION OF THE HOLOTYPE: An adult female (ROM 26381), head width 74% of length, length 46% of SVL; snout short, rounded in dorsal view, rounded in profile, protruding beyond margin of lower jaw; eye very large, prominent, 72% of snout; eyelid broader than interorbital distance. Top of head flat; canthus rostralis rounded; loreal region vertical, concave; lip flared just anterior to orbit; nostril about three­fourths distance from eye to tip of snout; supratympanic fold indistinct, slight swelling above tympanum; tympanum round, distinctly visible, separated from eye by distance equal to that of TMP, 41% of EYE. Choanae ovoid; vomerine dentigerous processes absent. Tongue cordiform, distinctly notched posteriorly, free for approximately two­thirds its length.

Forearms robust; fingers moderately short, slender, hand 28% of SVL, relative lengths of fingers I <II <IV <III, ventromedial callous pad on fingers II and III for twothirds length of finger; disks greatly expanded (>2× base of phalanges), relative pad size II <I <IV <III, pad length (finger III) equal to pad width, ventral circummarginal grooves present; terminal phalanges T­ shaped; subarticular tubercles conical. Hindlimbs moderately robust; tibia length 60% of SVL; foot length 84% of SVL; relative toe lengths I <II <III <V <IV; inner tarsal fold absent; feet fully webbed to disks, but as a fringe from distal subarticular tubercle of IV, lateral fringe on toe V to terminal phalanx; toes long, slender, with enlarged disks, smaller than those on fingers, relative pad size I = II = III> IV k V, pad width (IV) 83% of length, each pad with ventral circummarginal grooves; subarticular tubercles prominent and conical; inner metatarsal tubercle ovoid, long; outer metatarsal tubercle absent.

Xiphisternum narrow, notched posteriorly.

Skin on dorsum smooth, dorsolateral folds prominent in form of granules; small tubercles posteroventrally to tympanum, distinct granules on flanks and dorsum to cloaca; cloacal opening unmodified, directed posteriorly at upper level of thighs.

COLOR IN LIFE (in preservative): Dorsum green (livid blue), flanks brown (brown) and green (gray), each a prominent white spot; granules of dorsolateral fold golden (white); lip­stripe white (silvery white) from nostril to above insertion of arm; tympanum dark brown; loreal region dark brown (black); dorsal surfaces of limbs mottled brown and yellow with indistinct dark brown (black) transverse bands; posterior surface of thighs yellow with brown (black) mottling; webbing marbled white (translucent) and dark brown; venter creamy white; iris golden, pupil outlined in a striking yellow and red border.

SECONDARY SEXUAL CHARACTERS: Gravid females have immaculate white eggs. They are approximately 1.5 times larger than males. Males have a proportionally smaller tympanum than females (TMP:EYE for males 0.29, for females 0.45). The EYE:SNT is also greater in females (0.72) than it is in males (0.51). Males have velvety nuptial pads extending across thumb, and paired gular pouches located at the angles of the jaw. Pectoral spines are absent.

MEASUREMENTS OF HOLOTYPE (in mm): SVL 55.7; SNT 7.8; HDL 25.4; HDW 18.8; EYE 5.6; IOD 5.6; TMP 2.3; TEY 2.4; HND 15.5; FGR 13.9; FPL 2.8; FPW 2.8; TIB 33.6; FTL 47.0; TPL 2.3; TPW 1.9.

VARIATION OF PARATYPES: The loreal region on some specimens varies from dark brown to green. The large white spot on the flanks is sometimes accompanied by smaller ones. Flanks also have varying degrees of white mottling. The venter of some specimens has light mottling on the chest and chin. Variation in all type material is given in table 15.

MEASUREMENTS OF FEMALE PARATYPES (in mm, n = 8, ROM 38500, 38503, 38507, 38512, 38516, 38517, 38526, 38530): SVL 55.0 ± 1.2 (53.3–57.6); SNT 7.3 ± 0.5 (6.8– 8.3); HDL 17.8 ± 1.6 (16.7–19.4); HDW 17.2 ± 0.6 (15.6–17.6); EYE 5.8 ± 0.4 (5.3– 6.4); IOD 10.0 ± 1.7 (10.0–11.3); TMP 2.5 ± 0.2 (2.3–3.0); FPW 2.3 ± 0.3 (1.8–2.8); TIB 34.3 ± 1.3 (32.7–36.4); TPW 2.3 ± 0.3 (1.8–2.8).

MEASUREMENTS OF MALE PARATYPES (in mm, n = 7, ROM 26383, 26386, 26387, 26389, 26390, 26392, 26394): SVL 36.2 ± 1.2 (34.8–38.1); SNT 4.5 ± 0.4 (4.0–4.9); HDL 18.4 ± 1.7 (16.5–21.0); HDW 12.3 ± 0.6 (11.1–13.0); EYE 4.2 ± 0.6 (3.2–5.2); IOD 3 ± 0.4 (2.3–3.6); TMP 1.1 ± 0.2 (1.0– 1.9); TEY 0.6 ± 0.2 (0.2–1.0); HND 10.1 ± 0.6 (9.0–11.1); FGR 9.0 ± 0.3 (8.5–9.6); FPL 1.7 ± 0.3 (1.2–2.0); FPW 2.0 ± 0.2 (1.7–2.2); TIB 22.1 ± 1.3 (19.1–23.5); FTL 29.7 ± 4.4 (27.1–40.4); TPL 1.6 ± 0.2 (1.2– 1.7); TPW 1.4 ± 0.1 (1.2–1.6).

ETYMOLOGY: The specific name is a patronym for the –Dao people (pronounced ‘‘zao’’) of northern Vietnam.

DISTRIBUTION AND ECOLOGY: This species is known from the vicinity of Sa Pa village, Lao Cai Province in northern Vietnam. The photograph of a froglet of R. livida (in Karsen et al., 1998) also documents the occurrence of R. daorum in Hong Kong. The distinctive granular dorsolateral fold of R. daorum is clearly visible in the misidentified frog.

In early May, male Vietnamese R. daorum are actively calling by 1000 hours on partially submerged rocks in cascades as well as in vegetation adjacent to the streams. Females, although not as common, can be found slightly farther away from streams in more dense vegetation. One male (ROM 26394) was found in amplexus with the holotype (fig. 6E).

REMARKS: Rana daorum differs substantially from R. graminea ( Boulenger, 1899; Bourret, 1942) despite having a dorsolateral fold. Its small, forked xiphisternum differs from the large, deeply notched element of other members in the Rana chloronota complex and the subgenus Odorrana (sensu Fei et al., 1990) and more closely resembles that of male Huia nasica ( Yang, 1991b) . Huia nasica shares range, habitat and morphological similarities with the Rana chloronota complex: greatly expanded finger and toe disks, tremendous sexual dimorphism in size, white eggs, paired gular pouches, and a high chirplike call ( Boulenger, 1920; Pope, 1931; Bourret, 1942).

Figures

Fig. 11. Members of the Rana chloronota complex. (A) Male and female and (B) male R. chloronota, Nghe An Province, Vietnam. C. R. graminea, male, Hainan Island, China, (NLO field series 26375). D. R. bacboensis, new species, holotype, a female from Nghe An Province, Vietnam. (E) Female holotype and male paratype in amplexus, and (F) juvenile paratype of R. daorum, new species, Lao Cai Province, Vietnam.

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.