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Colostethus kuara sp. nov. Figures 5 C and 16 Suggested common names. golden-bellied rocket frog (English) / rana cohete de vientre dorado (Spanish). Phyllobates latinasus – Dunn (1924: 6) Phyllobates kingsburyi (part) – Dunn (1931: 391) Phyllobates pratti (part) – Dunn (1957: 78) Colostethus latinasus – Santos et al. (2009: fig. S 3, table SS 3), Pyron and Wiens (2011: 576) Colostethus pratti (part) – Samudio et al. (2015: 11), Pyron and Wiens (2011: 576), Savage (1968: 757), Edwards (1971: 148), Villa et al. (1988: 13), Jaramillo et al. (2010: 617, 633), Köhler (2011: 130, 131) Colostethus pratti - like – Grant et al. (2006: 132, 133, 177, 181, 233, 243, 250), Grant et al. (2017: S 38, S 40 – S 41, S 60) Colostethus aff. pratti. – Ibáñez et al. (2017: 93–96) Holotype. MHCH 2705, adult female, collected by AB and Konrad Mebert on 12 December 2012 in camp 2, Serranía de Pirre, Rancho Frío field station, El Real de Santamaría, Pinogana District, Darién Province, Parque Nacional Darién (PND), Republic of Panama (7.9595, –77.7037, 1230 m). Paratypes. SMF 97185 male from Pavarandó, camp 2, Jingurudó, Sambú District, Emberá – Wounaan (7.7587, –78.0916, 670 m), collected by AB and MV on 15 November 2012. SMF 97171 female and AB -1203 juvenile, same collecting data as holotype; SMF 97165 from Cana River, Cana field station, Chimenea trail, Boca de Cupe, Pinogana District, Darién Province, PND, Republic of Panama (7.7560, –77.6857, 530 m), collected by AB and Yorlis Cáceres on 9 September 2011. SMF 97164, female from Pirre ridge, El Real, Pinogana District, Darién Province, PND, Republic of Panama (7.9774, –77.7085, 1120 m), collected by AB, Yorlis Cáceres and Juan Pablo Rodríguez on 11 August 2011. MHCH 2687 male from camp 3, Púcuro River, Púcuro, Pinogana District, Darién Province, PND, Republic of Panama (8.0350, –77.3472, 620 m), collected by AB on 7 June 2012. SMF 97169 male from camp 4, above Púcuro River, Púcuro, Pinogana District, Darién Province, PND, Republic of Panama (8.0575, –77.3702, 1040 m), collected by AB on 8 June 2012. Diagnosis. Colostethus kuara 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 crests absent; (5) vocal sac well developed, vocal slits present, situated under the center of the orbit level, no nuptial pads; (6) appressed finger I, II and IV reaching distal subarticular tubercle of finger III; disk pads even; (7) no fringes or webbing on fingers; (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) toes weakly webbed according to the following formula: II 2 + – 3 + III 3 + – 4 + IV 4 + – 2 + V; (11) subarticular tubercles present (one on toes I and II, two on toes III and V, three on toes IV); inner metatarsal tubercle elongated along longitudinal axis, outer metatarsal tubercle slightly 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; dorsal ground coloration of thighs brown, grey or orangish, uniform or with dark transversal stripes; posterior color of thighs the same as dorsal color, with dark longitudinal band, with or without a yellow longitudinal stripe beneath; venter translucent, white, cream or yellow, with yellow coloration at the lower parts near groins; groins with yellow coloration, axillae with yellow coloration present or absent; ventral coloration of thighs and calves yellow; forelimbs with the same coloration, with or without transversal dark bands, humeri with or without longitudinal dark bands anteriorly and posteriorly; (13) axilla – groin stripe long, exceeds a mid-distance anteriorly from groin without interruption; at least lower one third of a tympanum is white; (14) SVL in males (n = 6): 17.9 ± 1.28 (16.2–20.1) mm, females (n = 6): 20.1 ± 1.67 (17.1–21.6) mm; (15) advertisement calls are performed in a chorus, at dominant frequency of 5242.2 ± 107.03 (5140–5353) Hz, with a note repetition rate of 12.8 ± 0.88 (12.1–13.8) notes / s and with a number of notes within one call being up to 49. Comparisons. Colostethus kuara sp. nov. differs from C. panamansis in maximum SVL of adult males (17.9 mm vs. 27 mm in C. panamansis), in degree of toe webbing (weak vs. moderate in C. panamansis) and in the coloration of throat (conspicuously mottled or reticulated brown or grey in C. panamansis females). Males’ advertisement calls consist of bouts of 26–49 notes (vs. 2–5 notes per call in C. panamansis), single notes have a duration of 26.3 ± 5.57 ms (vs. 60 ms in C. panamansis). Colostethus kuara 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) and in size (C. latinasus with male maximum SVL of 32.4 mm vs. 17.9 mm). Male advertisement calls differ significantly by its frequency (dominant frequency of 5140–5353 Hz vs. 2658–2673 Hz in C. latinasus). From Colostethus pratti, C. kuara sp. nov. differs by yellow coloration in groins and / or axillae (vs. white ventrolateral coloration in C. pratti); throat of C. pratti males is mottled with grey to dark brown (vs. uniform grey, white or yellowish in C. kuara sp. nov.); lower margin of the lateral dark stripe is interrupted by eye and tympanic annulus (unlike in C. pratti, where it goes below the tympanic annulus and no more than 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. kuara sp. nov.; no toe webbing (vs. toes weakly webbed in C. kuara sp. nov.); advertisement calls having a lower note repetition rate (12.8 ± 0.88 notes / s) than C. pratti (19 notes / s). Only female nurse frogs are known in C. pratti, whereas we collected a male nurse frog of C. kuara sp. nov. Colostethus kuara sp. nov. has a significantly shorter SVL in males than C. bocatoro sp. nov. (16.2–20.1 mm, 17.9 ± 1.28 mm vs. 19.0– 20.6 mm, 19.7 ± 0.83 mm; W = 3, p = 0.0303); a longer interorbital distance (t = 17.296, df = 9, p <0.001) and a shorter forearm length than C. bocatoro sp. nov. (t = – 2.4354, df = 9, p = 0.0377; a linear residual of SVL and either IO or FAL); with yellow coloration in groin and axilla (vs. no yellow coloration in C. bocatoro sp. nov.). It differs in finger webbing (absent vs. moderate in C. bocatoro sp. nov.) and in appearance of a tarsal keel (well defined vs. almost indiscernible in C. bocatoro sp. nov.). Advertisement calls are significantly faster (12.8 ± 0.88 notes / s; t = 8.4451, df = 7, p <0.001) than those of C. bocatoro sp. nov. (9.5 ± 0.35 notes / s) and consist of more notes (35.3 ± 12.10 notes, 26–49 notes) than the 11.2 ± 1.17 (11–13) notes of C. bocatoro sp. nov. calls. Colostethus kuara sp. nov. differs from C. donosensis sp. nov. by presence of a yellow coloration in groin and axilla (vs. only a ground color in C. donosensis sp. nov.). Advertisement calls consist of up to 49 notes (vs. up to 33 notes in C. donosensis sp. nov.) and have a dominant frequency of 5140–5354 Hz, 5242.2 ± 107.03 Hz, which is higher than in C. donosensis sp. nov. (4393–4759 Hz, 4585.7 ± 183.83 Hz), though it is correlated with the SVL. Colostethus kuara sp. nov. has a significantly shorter SVL in males than C. portobelensis sp. nov. (16.2–20.1 mm, 17.9 ± 1.28 mm vs. 20.3–22.0 mm, 21.1 ± 0.58 mm; t = – 5.7128, df = 10, p <0.001). The interorbital distance (IO) is significantly longer (t = 9.1513, df = 10, p <0.001); head size and forearm length are significantly smaller (HW: W = 0, p = 0.0022; and HL: t = – 4.2835, df = 10, p = 0.0016; FAL: t = – 3.9631, df = 10, p <0.01). In all the four parameters linear residuals of given distance and SVL were used. Colostethus kuara 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.) and by no finger webbing (vs. fingers weakly webbed in C. portobelensis sp. nov.). Number of notes per call is 35.3 ± 12.10 (26–49) notes is higher than the 10.7 ± 3.04 (6–16) notes of C. portobelensis sp. nov. calls. Note repetition rate of 12.8 ± 0.88 notes / s is also significantly faster (t = 14.629, df = 2.1095, p = 0.0037) than 5.3 ± 0.22 notes / s in C. portobelensis sp. nov. Single notes are significantly shorter (26.3 ± 5.57 ms; t = – 9.8288, df = 8, p <0.001) vs. 61.8 ± 5.12 ms in C. portobelensis sp. nov. Colostethus kuara sp. nov. has a shorter SVL in males than C. sanblasensis sp. nov. (16.2–20.1 mm, 17.9 ± 1.28 mm vs. 17.9–18.4 mm, 18.2 ± 0.25 mm); it has a uniform brown / grey dorsum or marbled with dark blotches with or without barely distinct median dark line (vs. distinct thin median dark line on uniform or slightly marbled brown / grey dorsum in C. sanblasensis sp. nov.); no finger webbing (vs. vestigial webbing between fingers III and IV in C. sanblasensis sp. nov.); toes weakly, but well discernible webbed (vs. only vestigial, barely discernible webbing in C. sanblasensis sp. nov.); male calls have a dominant frequency of 5140–5354 Hz, 5242.2 ± 107.03 Hz, that is higher than in C. sanblasensis sp. nov. (4336–4758 Hz, 4584.1 ± 171.03 Hz), though it is correlated with the SVL; while C. kuara sp. nov. was recorded only in a primary premontane / montane rainforest (530–1230 m), C. sanblasensis sp. nov. inhabits only in lower elevations
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