Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species Author BAIN, RAOUL H. Author LATHROP, AMY Author MURPHY, ROBERT W. Author ORLOV, NIKOLAI L. Author CUC, HO THU text American Museum Novitates 2003 2003-10-29 3417 1 60 http://www.bioone.org/doi/abs/10.1206/0003-0082%282003%29417%3C0001%3ACSOACF%3E2.0.CO%3B2 journal article 3413 10.1206/0003-0082(2003)417<0001:CSOACF>2.0.CO;2 0464179d-9549-4885-9ddd-32ffef32942e 0003-0082 4734880 Rana hmongorum , new species (Previously referred to as species 3, ‘‘Speckled’’) Figures 14A , 12I, J , 13D HOLOTYPE : (ROM field no. 19055) ROM 26376 an adult female approximately 5 km NW of Sa Pa village , near O Qui Ho Pass , Lao Cai Province , Vietnam (22°20̍09̎N, 103°50̍14̎E, elevation 1400 m ) collected 30 April 1995 by A. Lathrop and B. Hubley between 1900 and 2100 hours. The holotype had leg and liver tissue removed shortly after it was euthanised . PARATYPES : ROM 26370–26375, 26377– 26380, 39235, and 39236 collected in the vicinity of the holotype between 9 and 12 May 1995 by A. Lathrop and B. Hubley . ROM 26380, 39235, and 39236 are males and the remainder females . Additional specimens from the vicinity of the type locality include AMNH A­161480 collected on 27 August 1997 by D.R. Frost and C.J. Raxworthy , and ROM 39867–39896 (22°20̍58̎N, 103°46̍14̎E, elevation 1900 m ) collected between 30 April and 2 May 1999 by R.O. de Sa´ , C.T. Ho , A. Lathrop , T. Mason , R.W. Murphy , and N.L. Orlov. Male specimens are ROM 39874–39879, 39888, and 39890–39894; juveniles and subadults ROM 39867, 39868, 39880, 39895, and 39896; all others are females . DIAGNOSIS: Rana hmongorum , 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 59 mm ( 54–65 mm ), females 80 mm ( 74– 87 mm ); (3) vomerine teeth in rows posteromedial to choanae; (4) lip­stripe yellow, indistinct, extending across upper lip, terminating in a row of glandules; (5) head broad, bluntly rounded in profile; (6) tympanum round, distinct, TMP:EYE in males (0.43) greater than females (0.31); (7) supratympanic fold weak; (8) dorsal skin smooth with pustules becoming more pronounced laterally, pustules may form two dorsolateral lines resembling a fold, ventrum smooth; (9) dorsum and forearms green with black spots, flanks reddish brown with yellowish pustules, transverse bars on fore– and hindlimbs; (10) median callous pad on fingers II and III to proximal tubercle; (11) disks on fingers and toes moderately expanded (<2× base of phalanges); (12) feet fully webbed to disks, but as a fringe from distal subarticular tubercle of IV, lateral fringes on toes I and V to terminal phalanges, webbing brown­gray; (13) subarticular tubercles distinct, conical; inner metatarsal tubercle ovoid; (14) terminal phalanges T­ shaped; (15) xiphisternum large, deeply notched posteriorly; (16) males with nuptial pad; gular pouches and pectoral spines absent; (17) eggs white. COMPARISONS: Rana hmongorum superficially 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. hosii , R. jingdongensis , R. junlianensis , R. kwangwuensis , R. leporipes , R. livida , R. margaretae , R. schmackeri , R. sinica , and R. tiannensis (table 12). The white eggs of R. hmongorum distinguish it from H. nasica , Rana andersonii , R. chalconota , R. grahami , R. junlianensis , R. margaretae , R. schmackeri (all with white eggs with a melanic animal pole), and R. bacboensis (fully pigmented eggs). The dorsum and flanks with distinct pustules distinguishes R. hmongorum from all of the above species, except R. jingdongensis , and R. grahami ; R. daorum has two distinct rows of granular dorsolateral folds. Absence of gular pouches further differentiates it from H. nasica , R. archotaphus , R. bacboensis , R. chloronota , R. daorum , R. graminea , R. junlianensis , R. kwangwuensis , R. schmackeri , and R. tiannensis . Rana hmongorum has an indistinct yellow lipstripe distinguishing it from all other cascade ranids above, except R. chalconota , R. jingdongensis , R. junlianensis , and R. grahami . The presence of webbing to the base of the toe pad distinguishes R. hmongorum from R. leporipes (webbing to distal phalanx) and R. archotaphus . Larger females and the absence of an outer metatarsal tubercle further distinguish R. hmongorum from R. archotaphus (female SVL 59–62 mm ) and R. chalconota (SVL 46–59 mm ). Absence of ventral spines in males differentiates R. hmongorum from R. andersonii , R. grahami , R. jingdongensis , R. junlianensis , R. margaretae , and R. schmackeri . Rana sinica is further differentiated from R. hmongorum by its indistinct, skin­covered tympanum, the relative length of its digits (I <II <IV for R. sinica , II <I <IV for R. hmongorum ), and its rounded distal phalanges (T­ shaped in R. hmongorum ). Huia nasica is different in that it has a smooth, olive­brown dorsum, a longer head and smaller size ( 67 mm vs.> 75 mm for females, 44–49 mm vs. 54–59 for males). The small male of R. hmongorum further distinguishes it from R. jingdongensis , R. junlianensis , R. andersonii , and R. grahami (SVL 62–81 mm for R. jingdongensis , 70– 80 mm for R. andersonii , 66–84 mm for R. grahami ). The relative length of fingers further distinguishes it from R. hainanensis (II <IV <I <III) and R. hejiangensis (II <I <III <IV). 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 . DESCRIPTION OF THE HOLOTYPE : ROM 26376, an adult female, head length greater than width (136%), head width 36% of SVL, length 49% of SVL; snout short, protruding beyond margin of lower jaw, rounded in dorsal view, bluntly rounded in profile; eye large, prominent, 88% 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 indistinct, slight swelling above the tympanum; tympanum round, distinctly visible, separated from eye by distance equal to TMP, 26% of EYE. Choanae ovoid; vomerine dentigerous processes prominent, posteromedial to choanae, each bearing numerous teeth. Tongue cordiform, distinctly notched posteriorly, free for approximately one­half its length. Forearms robust; fingers relatively short, slender, hands 25% of SVL, relative lengths of fingers II <I <IV <III; ventromedial callous pad on fingers II and III to proximal tubercle, disks moderately expanded (<2× base of phalanges), relative pad size III> IV> I> II, pad length (III) 85% of pad width; ventral circummarginal grooves on disks present; terminal phalanges T­ shaped; subarticular tubercles conical. Hindlimbs moderately robust; tibia length 57% of SVL; foot length 40% 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 fringes on toes I and V to terminal phalanges; toes long, slender, with large, triangularly rounded disks, relative pad size I = II = III> IV k V, pad width (IV) 78% of pad length, ventral circummarginal grooves present; subarticular tubercles prominent and conical; inner metatarsal tubercle ovoid, long; outer metatarsal tubercle absent. Xiphisternum large, deeply notched posteriorly. Skin on dorsum smooth with large pustules, particularly on flanks, sacrum, and around cloaca; cloacal opening unmodified, directed posteriorly at upper level of thighs. COLOR IN LIFE (in preservative): Dorsum green (dark brown) and reddish brown (dark brown) with black spots. Sides reddish brown (gray) and yellow (cream), with some black spotting. Pustules on flanks yellowish white (creamy white), with black around the bases. Cloacal region dark brown (black). Lip­stripe creamy white (gray); dorsal surface of forearms with green (gray) patch; limbs mottled yellow (light brown) and brown (dark brown) with black transverse bands; anterior and posterior surfaces of thighs with brown (dark brown) marbling on yellow (cream); webbing uniformly brown; venter creamy white (creamy yellow). Iris greenish yellow or brown, pupil outlined with a yellow border. SECONDARY SEXUAL CHARACTERISTICS: Holotype is a gravid female with white eggs, 2 mm diameter. Adult female SVL 135% that of males. Males have slightly larger tympanum than do females, and velvety nuptial pads on thumb. Paired gular pouches and pectoral spines absent. Venter of males varies from immaculate creamy white to darkly mottled brown (gray in preservative). MEASUREMENTS OF HOLOTYPE (in mm): SVL 86.8; SNT 11.1; HDL 42.3; HDW 31.1; EYE 9.7; IOD 6.4; TMP 2.5; TEY 3.9; HND 22.3; FGR 16.5; FPL 2.3; FPW 2.7; TIB 49.2; FTL 35.0; TPL 3.0; TPW 2.3. VARIATION OF PARATYPES : Pustules can sometimes form dorsolateral lines, which superficially resemble folds. Pustules can be white. Variation in all type material is given in table 15. MEASUREMENTS OF FEMALE PARATYPES (in mm, n = 9; ROM 26370–26375, 26377– 26379): SVL 80.2 ± 4.3 (74.3–86.8); SNT 11.0 ± 0.5 (10.1–11.7); HDL 38.8 ± 1.9 (37.0–41.0); HDW 29.8 ± 1.0 (29.2–31.3); EYE 9.3 ± 1.1 (7.6–10.8); IOD 7.0 ± 0.3 (6.4–7.3); TMP 2.9 ± 0.4 (2.5–3.3); TEY 3.0 ± 0.6 (2.1–3.9); HND 23.2 ± 1.3 (22.1– 25.6); FGR 18.6 ± 1.1 (16.5–19.6); FPL 3.0 ± 0.4 (2.3–3.4); FPW 2.6 ± 0.4 (2.0–3.1); TIB 47.6 ± 2.3 (43.1–49.2); FTL 39.6 ± 5.0 (35.0–48.3) TPL 2.8 ± 0.6 (2.0–3.5); TPW 2.4 ± 0.5 (1.9–3.3). MEASUREMENTS OF MALE PARATYPES (in mm, n = 12; ROM 26380, 30876, 39235, 39236, 39874, 39875, 39877, 39879, 39888, 39891, 39892; AMNH A­161480): SVL 59.4 ± 3.8 (54.7–65.3); SNT 9.4 ± 0.9 (8.7– 10.4); HDL 24.0 ± 3.0 (20.7–30.4); HDW 20.0 ± 1.2 (18.9–21.1); EYE 7.3 ± 0.5 (6.7– 8.3); IOD 5.3 ± 0.7 (4.3–6.5); TMP 3.0 ± 0.4 (2.4–3.7); TEY 2.5 ± 0.3 (2.2–3.0); HND 17.2 ± 1.4 (14.1–19.4); FGR 13.3 ± 1.2 (12.8–14.8); FPL 2.5 ± 0.5 (1.6–3.6); FPW 2.1 ± 0.6 (1.0–2.4); TIB 38.6 ± 4.5 (33.4–51.2); FTL 47.2 ± 5.7 (30.4–52.1); TPL 2.3 ± 0.4 (1.7–2.9); TPW 1.7 ± 0.4 (0.9–2.2). ETYMOLOGY: The specific name is a patronym for the Hmong (pronounced ‘‘huhmung’’) people, an ethnic group in the northern montane regions of Vietnam . Their assistance made it possible for us to document the fauna of Hoang Lien Mountains. DISTRIBUTION AND ECOLOGY: This species is only known only from the vicinity of Mount Fan Si Pan at approximately 1900 m or above. Adults were found along the cascades of a waterfall and along the mossy slopes of a human­made culvert.