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Colostethus sanblasensis sp. nov. Figures 5 E and 18 Suggested common names. San Blas rocket frog (English) / rana cohete de San Blas (Spanish) Colostethus pratti (part) – Savage (1968: 757), Edwards (1971: 148), Villa et al. (1988: 13), Jaramillo et al. (2010: 617, 633), Köhler (2011: 130, 131) Holotype. MHCH 2708, adult male, collected by AB and MV on 6 October 2012, in Nurra (9.0296, –78.0360, 320 m), Comarca Guna de Wargandí, Republic of Panama. Paratypes. SMF 97197 male and SMF 97196 female, same collection data as holotype; SMF 97182 male and MHCH 2707 female from Bajo Pequeño, camp 3 Pechito Parao, Lajas Blancas, Cémaco District, Comarca Emberá – Wounaan, Republic of Panama (8.4800, –77.5194, 470 m), collected by AB and MV on 6 November 2012. Diagnosis. Colostethus sanblasensis sp. nov. is characterized by the following combination of characters: (1) dorsal skin smooth anteriorly, with scattered tubercles in the last 1 / 3 of the body, ventral skin texture smooth, usually with disperse warts present around the cloaca; (2) tympanic annulus present, but only the lower part is clearly visible, tympanic membrane absent; (3) snout truncate in dorsal view and rounded in profile; (4) rounded supraocular tubercle barely discernible, cranial crest absent; (5) vocal sac well developed, vocal slits present, situated under the center of the orbit level, no nuptial pads; (6) appressed finger II and IV reaching distal subarticular tubercle of finger III, while finger I not; disk pads even; (7) no fringes on fingers, vestigial webbing between fingers III and IV; (8) palmar tubercle rounded, flattened and larger than thenar tubercle; thenar tubercle low and elongated; subarticular tubercles rounded and globular, first tubercle more evident; supernumerary tubercles barely evident; (9) heel smooth; (10) vestigial webbing between toes II – IV; (11) subarticular tubercles present (one on toes I and II, two on toes III and V, three on toe IV); inner metatarsal tubercle elongated along longitudinal axis, outer metatarsal tubercle rounded and smaller than the inner; tarsal keel well defined; (12) dorsal ground color of head and trunk in life brown or grey, uniform or with dark blotches, with a thin median dark line continuously leading from snout to the base of the back or disrupted but with remnants still clearly visible; dorsal ground coloration of thighs reddish or grey, uniform or with dark transversal stripes; posterior color of thighs reddish or grey with dark longitudinal band on thighs (if grey, then with a yellow longitudinal stripe beneath the dark longitudinal band); venter translucent, suffused with cream or yellow, with yellow coloration at the lower parts near groins; groins with reddish or yellow coloration at the femoral part and yellow coloration at the abdominal part; ventral coloration of thighs white with reddish margins (if reddish thighs dorsally) or yellow (if grey thighs dorsally); ventral coloration of calves uniform reddish or white with reddish margins (if thighs are reddish) or uniform yellow (if thighs are grey / yellow); forelimbs reddish dorsally and white ventrally (if hind limbs reddish), or reddish or grey dorsally and yellow ventrally (if hind limbs grey / yellow); (13) axilla – groin stripe short, interrupted, extending from the groin anteriorly to about midway toward the axilla; (14) SVL in males (n = 4): 18.2 ± 0.25 (17.9–18.4) mm, females (n = 1): 15.3 mm; (15) advertisement calls are performed in a chorus, at dominant frequency of 4584.1 ± 171.03 (4336–4758) Hz, with a note repetition rate of 11.7 ± 1.98 (7.7–13.5) notes / s and with a number of notes within one call being up to 49. Comparisons. Colostethus sanblasensis sp. nov. differs from C. panamansis in maximum SVL of adult males (18.4 mm vs. 27 mm in C. panamansis) and in degree of toe webbing (toes moderately webbed in C. panamansis). Males’ advertisement calls consist of bouts of 34.3 ± 9.46 (22–49) notes (vs. 2–5 notes per call in C. panamansis), single notes have a duration of 31.4 ± 6.70 (22.3–39.3) ms (vs. 60 ms in C. panamansis). Colostethus sanblasensis sp. nov. differs from C. latinasus in male ventral coloration, throat and chest brown with white spots forming mottled pattern, darker in adult males, lighter but still conspicuous in adult females of C. latinasus, toes weakly webbed (vs. vestigial, barely visible webbing in C. sanblasensis sp. nov.) and in size, with C. latinasus male maximum SVL of 32.4 mm vs. 18.4 mm in C. sanblasensis sp. nov. males. Male advertisement calls differ in dominant frequency (2658–2673 Hz in C. latinasus vs. 4336–4758 Hz in C. sanblasensis sp. nov.). Colostethus pratti and C. sanblasensis sp. nov. differ by dorsal skin color, which is brown or gray with or without darker speckles (unlike in C. pratti it is uniform brown); C. sanblasensis sp. nov. has yellow or reddish coloration in groin and axilla (white ventrolateral coloration in C. pratti); throat of C. pratti males is mottled with grey to dark brown (vs. uniform white or yellowish in C. sanblasensis sp. nov.); the lower margin of the lateral dark stripe is interrupted by eye and tympanic annulus (unlike in C. pratti, where the stripe goes below the tympanic annulus and the eye or only about one half of the eye length reaches out of the dark stripe). In C. pratti the lateral dark stripe seems to be relatively thicker than in C. sanblasensis sp. nov.; advertisement calls of C. pratti males consist of shorter notes (20 ms vs. 31.4 ± 6.7 ms) with a higher rate (19 notes / s vs. 11.7 ± 1.98 notes / s). Only female nurse frogs are known in C. pratti, whereas we collected a male nurse frog of C. sanblasensis sp. nov. Colostethus sanblasensis sp. nov. has a significantly smaller SVL in males than C. bocatoro sp. nov. (17.9–18.4 mm vs. 19.0– 20.6 mm, 19.7 ± 0.83 mm; W = 0, p = 0.0159) and longer interorbital distance than C. bocatoro sp. nov. (a linear residual of SVL and IO, respectively; t = 26.192, df = 7, p <0.001). Colostethus sanblasensis sp. nov. has yellow or reddish coloration in groin and axilla (vs. no yellow or reddish coloration in flank part of groin and axilla); vestigial webbing between finger III and IV (vs. moderately webbed fingers in C. bocatoro sp. nov.); well defined tarsal keel (vs. almost indiscernible in C. bocatoro sp. nov.); calls are faster (11.7 ± 1.98 notes / s; t = 6.5562, df = 6.2047, p <0.001) than those of C. bocatoro sp. nov. (9.5 ± 0.35 notes / s) and consist of significantly more notes (34.3 ± 9.5 notes, 22–49 notes; t = 7.2036, df = 5.1901, p <0.001) than 11.2 ± 1.17 (11–13) of C. bocatoro sp. nov. Colostethus sanblasensis sp. nov. differs from C. donosensis sp. nov. by presence of reddish or yellow coloration in groin and axilla (vs. only a ground color); by only vestigial toe webbing (vs. weak, but well discernible in C. donosensis sp. nov.); by a significantly higher note repetition rate (11.7 ± 1.98 notes / s vs. 6.0 ± 0.17 notes / s; t = 10.593, df = 7, p <0.001) and by a calling in a chorus with other males (vs. individually or with only one response to another male). Colostethus sanblasensis sp. nov. has a thin median dark line on a brown / grey back and head (vs. no median line or barely distinct among dark blotches in C. kuara sp. nov.); vestigial webbing between fingers III and IV (vs. no finger webbing in C. kuara sp. nov.); toe webbing vestigial, barely discernible (vs. weak but well discernible in C. kuara sp. nov.); male calls have a dominant frequency of 4336–4758 Hz, 4584.1 ± 171.03 Hz, that is lower than in C. kuara sp. nov. (5140–5354 Hz, 5242.2 ± 107.03 Hz), though it is correlated with the SVL; while C. sanblasensis sp. nov. was recorded in elevations of 170–470 m, C. kuara sp. nov. only in a primary premontane / montane rainforest (530–1230 m). Colostethus sanblasensis sp. nov. has a significantly shorter SVL in males than C. portobelensis sp. nov. (17.9–18.4 mm vs. 20.3–22.0 mm, 21.1 ± 0.58 mm; t = – 9.6096, df = 8, p <0.001); a significantly longer interorbital (IO: t = 10.443, df = 8, p <0.001) and axilla – groin distance (AGD: t = 3.7698, df = 8, p <0.01); and a significantly smaller head (HW: W = 0, p = 0.0095; and HL: t = – 2.5167, df = 8, p = 0.0360) and forearm length (FAL: t = – 3.4957, df = 8, p <0.01). In all the five parameters linear residuals of given distance and SVL were used. Colostethus sanblasensis sp. nov. also differs from C. portobelensis sp. nov. by a belly and throat of the same color (vs. pale belly and darker throat – greyish or purplish – in C. portobelensis sp. nov.). Only vestigial webbing between fingers III and IV (vs. weakly webbed fingers in C. portobelensis sp. nov.). Number of notes 34.3 ± 9.5 (22–49) per call is significantly higher (t = 7.0255, df = 6.1008, p <0.001) than 10.7 ± 3.04 (6–16) of C. portobelensis sp. nov. Note repetition rate of 11.7 ± 1.98 notes / s is also significantly faster (t = 17.059, df = 5.4311, p <0.001) than 5.3 ± 0.22 notes / s (p <0.05) in C. portobelensis sp. nov. Single notes are significantly shorter (31.4 ± 6.7 ms; t = – 9.6526 df = 11, p <0.001) vs. 61.8 ± 5.12 ms in C. portobelensis sp. nov. Males of C. sanblasensis sp. nov. call in a chorus with other conspecific males (vs. individually or with only one response to another male). Colostethus sanblasensis sp. nov. has a significantly shorter SVL in males than C. chucantiensis sp. nov. (17.9–18.4 mm, 18.2 ± 0.25 mm vs. 18.4–25.1 mm, 21.8 ± 1.75 mm; t = 7.5917, df = 14.563, p <0.001); it has reddish or grey ground coloration of thighs dorsally (if grey, then with a yellow longitudinal stripe beneath the dark longitudinal band on thighs posteriorly) vs. uniform grey or brown dorsal coloration of thighs in C. chucantiensis sp. nov.; vestigial webbing between fingers III and IV (vs. no finger webbing in C. chucantiensis sp. nov.); toe webbing vestigial, barely discernible (vs. weak but well discernible in C. chucantiensis sp. nov.); males call in a chorus with up to 5 resp
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