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 banaorum , new species (Previously referred to as species 6, ‘‘Southern Big Eye’’) Figures 12M, N , 13F , 14E, F HOLOTYPE : (ROM field no. 7145) ROM 39944, an adult female from Tram Lap , An Khe District , Gia Lai Province , Vietnam (14°26̍39̎N, 108°32̍97̎E), elevation ca. 900 m, collected on 15 June 1996 by R.H. Bain, A. Lathrop, R.W. Murphy, and N.L. Orlov. The holotype had leg and liver tissue removed shortly after it was euthanised. TABLE 15 Variation in Body Proportions for Rana chloronota sensu stricto and Vietnamese Species of the Rana chloronota Complex Given as Means (in mm) ± 1 SD and ranges See Materials and Methods for abbreviations. PARATYPES : Four males (ROM 39912– 39913, 39915–39916) and eight females (ROM 39899–39901, 39928, 39929, 39931, 39936, and 39941) collected with holotype between 15 and 29 June 1996 by R.H. Bain , A. Lathrop , R.W. Murphy , and N.L. Orlov. ROM 39716–39720, 39920 –39922, and 39924–39926 (males); 39942 (subadult) from the Cha River , Buon Loi , An Khe District , Gia Lai Province , Vietnam (elevation ca. 900 m ), collected on 27 June 1996 by R.H. Bain and N.L. Orlov. ROM 39700, and 39702–39705 males from Krong Pa, An Khe District , Gia Lai Province , Vietnam (14°20̍29̎ N, 108°28̍46̎E, elevation 850 m ), collected on 13–25 August 1997 by C.T. Ho , A. Lathrop , L.A. Lowcock , R.W. Murphy , and N.L. Orlov. ROM 25084–25086, 25100, 25102, and 25103 (males) collected from Buon Loi , An Khe District , Gia Lai Province , Vietnam (elevation ca. 700–750 m ), 5 November 1993 by I.S. Darevsky and N.L. Orlov . DIAGNOSIS: Rana banaorum , 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 50 mm ( 42–55 mm ), females 93 mm ( 83–99 mm ); (3) vomerine teeth in rows oblique to choanae; (4) white lip­stripe extending across upper lip, terminating in glandule above insertion of arm; (5) head broad, bluntly rounded in profile; (6) tympanum round, distinct, very large, TMP:EYE in males (0.89) greater than females (0.75); (7) supratympanic fold weak; (8) dorsal skin shagreened, becoming granular laterally, with slight dorsolateral folds; (9) dorsum usually brown with yellow and black spots, sometimes green with black spots; legs brown, lightly banded; (10) median callous pad on fingers II, and III to proximal tubercle; (11) disks on fingers and toes greatly enlarged (>2× base of phalanges); (12) feet fully webbed to disks, except medial side of IV, where it reaches disk as a fringe, lateral fringes on I and V to terminal phalanges, webbing brown; (13) subarticular tubercles and inner metatarsal tubercle distinct, conical; (14) terminal phalanges T­ shaped; (15) xiphisternum large, deeply notched posteriorly; (16) males with nuptial pads, paired gular pouches, pectoral spines absent; (17) eggs white. COMPARISONS: Among Asian cascade ranids, R. banaorum is most similar to sympatric R. morafkai and particularly R. chloronota but it can be anatomically distinguished from both by its dorsolateral folds (table 12). Rana banaorum can further be distinguished from R. morafkai by its larger males (SVL 42–55 mm , versus 39–45 mm ). Some Rana banaorum males have indistinct, microscopic spinules on the dorsal surface of the leg extending to the feet, forming a ‘‘saw­tooth’’ formation along the lateral edge of toe V, which is present only to a slight degree or absent in R. chloronota . Its broad, rounded snout differs from the obtusely pointed snouts of H. nasica , R. andersonii , R. chalconota , and R. schmackeri and from the depressed snouts of R. graminea and R. margaretae . A white lip­stripe differentiates R. banaorum from R. andersonii , R. bacboensis , R. grahami , R. hainanensis , R. hmongorum , R. jingdongensis , R. junlianensis , R. margaretae , R. sinica , R. schmackeri , and R. tiannensis . The gular pouches of R. banaorum distinguish it from R. andersonii , R. chalconota , R. grahami , R. hainanensis , R. hmongorum , R. hosii , R. jingdongensis , R. junlianensis , R. kwangwuensis , and R. margaretae . The dorsolateral folds of R. banaorum distinguish it from H. nasica , R. andersonii , R. bacboensis , R. chloronota , R. hainanensis , R. hejiangensis , R. jingdongensis , R. junlianensis , R. kwangwuensis , R. morafkai , R. livida , R. schmackeri , R. sinica , and R. tiannensis ; the dorsolateral folds of R. daorum are composed of distinct white granules, and dorsolateral pustules sometimes form folds on R. grahami and R. hmongorum . The absence of an outer metatarsal tubercle immediately differentiates R. banaorum from R. archotaphus and R. chalconota . The presence of white eggs differentiates R. banaorum from R. bacboensis (black eggs), Huia 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. banaorum from R. andersonii , R. grahami , R. jingdongensis , R. junlianensis , R. margaretae , and R. schmackeri . Rana banaorum further differs from Huia nasica by its larger females ( 83–99 mm vs. 67 mm in H. nasica ). It is further differentiated from R. sinica by its uncovered, distinct tympanum (indistinct, covered by skin in R. sinica ), large disks (small in R. sinica ) and its relative finger lengths (I <II <IV for R. sinica , II <I <IV for R. banaorum ) and T­ shaped distal phalanges (rounded in R. sinica ). Rana leporipes further differs from R. banaorum with its white supratympanic fold (not colored in R. banaorum ), webbing to distal phalanges (webbing to disks in R. banaorum ), and T­ shaped distal phalanges (oblong, somewhat rounded in R. leporipes ). DESCRIPTION OF HOLOTYPE : ROM 39944, an adult female, head width 78% head length, length 46% SVL; snout short, acutely rounded in dorsal view, bluntly rounded in profile, protruding beyond margin of lower jaw; eye large, prominent, 50% 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 arm; tympanum very large, round, distinctly visible, separated from eye by distance equal to TMP, 80% of eye length. 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 23% SVL, relative finger lengths II <I <IV <III, ventromedial callous pad on fingers II and III to proximal tubercle, disks greatly expanded (>2× base of phalanges), relative pad size II <I <IV <III, pad length (III) 94% of pad width, ventral circummarginal grooves present; terminal phalanges T­ shaped; subarticular tubercles conical. Hindlimbs moderately robust; tibia length 65% SVL; relative toe lengths I <II <III <V <IV; inner tarsal fold absent; feet very large, 82% of SVL, fully webbed to terminal phalanges except the inner side of IV where the webbing reaches the disk as a fringe; lateral fringe on toes I and V to terminal phalanges; toes long, slender, with large, rounded triangular disks; relative pad size I = II = III> IV> V, pad length (IV) equals pad width; each pad 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, with small pustules on flanks; dorsolateral folds weak; small tubercles posteroventral to tympanum; prominent granules on flanks and around cloaca; cloacal opening unmodified, directed posteriorly, at upper level of thighs. COLOR IN LIFE (in preservative): Dark brown (olive­brown, beige), flanks gray with yellow spots (white spots); lip­stripe brown anteriorly and creamy white from level of the eye posteriorly (white); loreal dark brown (black); tympanum beige with dark brown central ring; iris golden brown; dorsal limbs grayish brown with black banding; cloacal region dark brown (black); webbing marbled white on dark brown (uniformly brown); venter creamy white, ventral surfaces of limbs creamy yellow with black mottling. SECONDARY SEXUAL CHARACTERS: The holotype is a gravid female with immaculate white eggs ( 2 mm in diameter). Gravid females have an SVL nearly twice that of males (mean SVL female, 93 mm ; male 50 mm ). Male TMP:EYE (0.89) is larger than females (0.75). Males have velvety nuptial pads on the thumb and paired gular pouches located at the angle of jaw. Pectoral spines are absent. MEASUREMENTS OF HOLOTYPE (in mm): SVL 98.0; SNT 13.5; HDL 45.4; HDW 35.6; EYE 6.8; IOD 7.4; TMP 5.4; TEY 4.7; HND 22.7; FGR 19.4; FPL 3.1; FPW 3.3; TIB 63.7; FTL 80.0; TPL 3.4; TPW 3.4. VARIATION OF PARATYPES : The skin on the dorsum varies from smooth to shagreened with small pustules on flanks. The lip­stripe varies from creamy yellow (white in alcohol) throughout its length to brown anteriorly and creamy white from the level of the eye posteriorly. The dorsal skin is variable from light green to dark brown (olive, brown, beige, or livid blue in alcohol) with or without large black spots. Variation in all type material is given in table 15. MEASUREMENTS OF FEMALE PARATYPES (in mm, n = 8, ROM 39899–39901, 39928, 39929, 39931, 39936, and 39941): SVL 92.7 ± 5.6 (83.4–98.7); SNT 13.8 ± 1.6 (12.4– 16.8); HDL 43.8 ± 3.2 (37.5–48.8); HDW 30.6 ± 1.5 (26.7–33.4); EYE 6.1 ± 0.7 (4.7– 6.9); IOD 8.2 ± 0.9 (6.8–10.1); TMP 4.5 ± 0.6 (3.5–5.6); TEY 4.5 ± 0.6 (3.5–5.2); HND 22.3 ± 2.8 (17.5–25.0); FGR 18.2 ± 2.8 (12.9–20.8); FPL 3.4 ± 0.4 (2.8–4.3); FPW 3.1 ± 0.2 (2.8–3.5); TIB 58.1 ± 3.2 (52.7–63.7); FTL 68.0 ± 7.0 (58.3–78.0) TPL 3.4 ± 0.6 (2.3–4.3); TPW 2.8 ± 0.4 (2.4–3.4). MEASUREMENTS OF MALE PARATYPES (in mm, n = 14, ROM 25084–25086, 25100, 25102, 25103, 39912, 39913, 39915, 39916, 39920–39922, 39924): SVL 50.5 ± 3.7 (42.5–54.6); SNT 7.4 ± 1.5 (3.1–8.7); HDL 26.1 ± 1.4 (24.6–28.1); HDW 17.4 ± 0.6 (17.8–18.3); EYE 4.0 ± 0.5 (3.1–5.1); IOD 3.8 ± 0.4 (3.4–4.7); TMP 3.5 ± 0.6 (2.0– 4.7); TEY 1.7 ± 0.6 (0.9–3.6); HND 14.0 ± 1.2 (11.8–16.0); FGR 11.1 ± 1.1 (9.2–12.7); FPL 1.9 ± 0.3 (1.4–2.6); FPW 1.6 ± 0.2 (1.4–2.0); TIB 30.6 ± 4.6 (24.1–43.6); FTL 32.6 ± 6.8 (23.3–42.7); TPL 1.9 ± 0.4 (1.3– 2.4); TPW 1.5 ± 0.3 (1.1–2.1). ETYMOLOGY: The specific name is a patronym for the Ba Na people, an ethnic group living on the Tay Nguyen Plateau (Central Highlands) of south­central Vietnam where this species occurs. DISTRIBUTION AND ECOLOGY: Rana banaorum is known only from the Tay Nguyen Plateau of the Central Highlands , Gia Lai Province , Vietnam . It inhabits forested montane river systems on or near rapids or waterfalls of primary and disturbed second growth. In May and June males have tight, leathery gular pouches, suggesting that they are not calling and that the breeding season occurs during some other time . REMARKS: Inger and Chanard (1997) and Inger et al. (1999) noted that specimens of Rana chloronota (as R. livida ) from An Khe have more pronounced dorsolateral folds than elsewhere in Vietnam . These frogs do not fit Bourret’s (1942) interpretation of R. graminea as a ‘‘northern variety’’ of R. chloronota . These differences likely reflect the occurrence of three sympatric species of the R. chloronota complex from this region. Some male R. banaorum have indistinct, microscopic spinules on the dorsal surface of the leg extending to the feet, forming a ‘‘saw­tooth’’ formation along the lateral edge of toe (seen in FMNH specimens that are not part of the type series).