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 livida ( Blyth, 1856 ) Figures 6E, F , 7E, F , 9 Rana livida ( Boulenger, 1887 ) NEOTYPE : BMNH 1889.3.25.48 an adult female from Thagata Juwa , Village on the hills southwest of Mt. Mooleyit , Dawna Mountains , Myanmar ( 400–500 m ), by M.L. Fea in early 1887. DIAGNOSIS: Rana livida , 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 females 89.4, 97.1 mm ; (3) vomerine teeth in rows oblique to choanae; (4) lip­stripe white, extending across upper lip, terminating in glandule above insertion of arm; (5) head broad, snout rounded in dorsal view, bluntly rounded in profile; (6) tympanum round, distinct, TMP:EYE 0.41; (7) supratympanic fold weak; (8) dorsal skin smooth to flanks, pustules dorsal to cloaca, dorsolateral folds absent, venter smooth; (9) dorsum uniform brown in preservative; forelimbs and hindlimbs lighter brown, without transverse bars; (10) median callous pad on finger III to proximal tubercle; (11) disks on fingers and toes enlarged (>2× base of phalanges); (12) feet fully webbed to disks, weak lateral fringes on I and V to terminal phalanges; (13) subarticular tubercles and inner metatarsal tubercle distinct, conical; (14) terminal phalanges T­ shaped; (15) xiphisternum large, deeply notched posteriorly; (17) eggs white. COMPARISONS: Rana livida superficially resembles other Asian cascade ranids, including Huia nasica , Rana andersonii , R. archotaphus , R. chalconota , R. chloronota , R. grahami , R. graminea , R. hainanensis , R. hejiangensis , R. hosii , R. jingdongensis , R. junlianensis , R. kwangwuensis , R. leporipes , R. margaretae , R. schmackeri , R. sinica , and R. tiannensis (table 12). The smooth, uniformly colored dorsum further distinguishes R. livida from all of the above species. The unpigmented eggs of R. livida distinguish it from H. nasica , Rana andersonii , R. chalconota , R. grahami , R. junlianensis , R. margaretae , and R. schmackeri , which are white with melanic poles. Huia nasica differs from R. livida in its longer, more pointed head, and smaller females (SVL 67 mm ). Rana hainanensis further differs from R. livida in relative lengths of fingers (II <IV <I <III for R. hainanensis , II <I <IV <III for R. livida ). Rana hejiangensis differs from R. livida in its relative finger size (II <I <III <IV) and smaller digital disks. Rana andersonii has a rough, olive­brown dorsum and small disks on fingers. Rana grahami has pustules on the dorsum and flanks (vs. smooth in R. livida ) and slightly swollen digital tips. Rana junlianensis has brown lip bands. Rana kwangwuensis has small digital disks. Rana schmackeri has a smooth, heavily spotted dorsum. Rana tiannensis has a rough brown dorsum with large, prominent lateral granulations. Rana chalconota differs from R. livida by having coarsely shagreened dorsum, distinct dorsolateral folds, pointed snout, smaller SVL (females 46–59 mm ), and an outer metatarsal tubercle. In R. archotaphus , webbing (toe IV) reaches base of distal subarticular tubercle ( R. livida , webbing to base of disk), females are smaller (SVL 59–62 mm ), and it has an outer metatarsal tubercle. Rana sinica differs from R. livida in lacking a lip­stripe, and in having smaller mature females ( R. sinica holotype 66.6 mm ), an indistinct, skin­covered tympanum (distinct and uncovered in R. livida ), nares halfway between the eye and tip of the snout (nearer the snout in R. livida ), and a different finger formula (I <II <IV for R. sinica , II <I <IV for R. livida ). Rana hosii differs from R. livida by its obtusely pointed head (rounded in R. livida ), dorsolateral folds, occasional bands on arms and legs, and feeble tarsal folds (dorsolateral and tarsal folds absent in R. livida ). Rana chloronota differs from R. livida with its solid­colored dorsum and dark sides (uniform in R. livida ), transverse bars on the limbs (absent in R. livida ), smooth dorsum, flanks granular (slightly granulose on the posterior thighs in R. livida ), and absence of white mottling on the flanks (present in R. livida ). Rana leporipes differs from R. livida in having smaller digital disks, a white supratympanic fold, weak dorsolateral fold, and webbing that only reaches the distal phalanges ( R. livida to base of the disk). Rana graminea further differs from R. livida with its nearly vertical (versus concave) loreal region and depressed head. DESCRIPTION: Head length greater than head width (103%), head width 35% of SVL, length 37% of SVL; snout short, protruding beyond margin of lower jaw, rounded in dorsal view, bluntly rounded in profile; eye large, prominent, 72% 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 weak, swollen rim of tissue dorsal to tympanum; tympanum round, distinctly visible, 44% of EYE. Choanae ovoid; vomerine dentigerous processes prominent, oblique, posteromedial to choanae, each bearing numerous teeth. Tongue cordiform, distinctly notched posteriorly, free for approximately one­third its length. Forearm robust; fingers moderately short, slender; hands 29% of SVL, relative lengths of fingers II <I <IV <III; ventromedial callous pad on III to proximal tubercle; disks greatly expanded (>2× base of phalanges), relative pad size II <I <IV <III; circummarginal grooves present ventrally; terminal phalanges T­ shaped; subarticular tubercles prominent, rounded. Hindlimbs moderately robust; tibia length 68% of SVL; FTL 85% of SVL; relative toe lengths I <II <III <V <IV; inner tarsal fold absent; feet fully webbed to base of toe disk, lateral fringes on I and V to terminal phalanges; toes long, slender, with large, obliquely rounded disks, relative pad size I = II = III> IV> V; ventral circummarginal grooves present; subarticular tubercles prominent, conical; inner metatarsal tubercle ovoid, long; outer metatarsal tubercle absent. Xiphisternum large, deeply notched posteriorly. Skin on dorsum smooth to flanks; dorsolateral fold absent; small tubercles posteroventral to tympanum; granules dorsal cloaca; cloacal opening unmodified, directed posteriorly, at upper level of thighs. COLOR IN PRESERVATIVE: Dorsum uniform dark brown to flanks, some white spots and mottling on flank; prominent lip­stripe white; tympanum brown; posterior surface of thighs light brown with white spots, no transverse banding; venter creamy white; ventral side of limbs creamy white; webbing dark gray. SECONDARY SEX CHARACTERS: BMNH 1889.3.25.47, an adult female bears white eggs. No male specimens were studied. MEASUREMENTS OF NEOTYPE (in mm): SVL 89.4; SNT 14.2; HDL 32.8; HDW 31.6; EYE 10.2; IOD 8.0; TMP 4.5; TEY 2.8; HND 25.5; FPL 3.2; TIB 60.8; FTL 76.2; TPL 2.6. MEASUREMENTS OF REFFERED SPECIMEN (BMNH 1889.3.25.47): SVL 97.1; SNT 16.7; HDL 36.1; HDW 33.4; EYE 11.9; IOD 9.2; TMP 4.6; TEY 4.1; HND 28.8; FPL 3.3; TIB 63.8; FTL 75.6; TPL 2.7. DISTRIBUTION AND ECOLOGY: The range of R. livida is unclear. The type material is lost and the locality information has been poorly recorded. Much confusion exists in its taxonomy. The only confirmed locality is that of the neotype and maps do not show Thagata Juwa village , as indicated in by Feae. Mount Mooleyit is currently found on maps as Mulayit Taung (16°N, 98°30̍E). The closest villages to Mulayit Taung are Kyeik­ywa, Daukkat­ywa, and Kyeik­don, to the west, and Mawkhi and Huthi, near the Thai border to the east. REMARKS: Male R. chloronota are substantially smaller than females and they posses paired subgular pouches and a larger tympanum. Presumably, male R. livida have the same attributes. Blyth (1852) appears to have included males in his original description; many of the specimens that Theobold assumed to be juveniles have whiter lip­stripes and a relatively large tympanum that is closer to the eye than in females. Although the color is not recorded in life, Blyth (1856) reported that R. livida is ‘‘uniform duskyplumbeous above, probably dull olive green when alive.’’ Boulenger (1920) recorded measurements of two R. livida females from Thagata, Tenasserim. It is not clear whether these two specimens are the same as those described above.