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Colostethus donosensis sp. nov. Figures 5 B and 15 Suggested common names. Donoso rocket frog (English) / rana cohete de Donoso (Spanish) Colostethus pratti (part) – Lips et al. (2006: 103, supplementary material), Brem and Lips (2008: 81, appendix 1), Crawford et al. (2010: 107), Savage (1968: 757), Edwards (1971: 148), Villa et al. (1988: 13), Santos et al. (2009: fig. S 3, table SS 3), Auth (1994: 4), Consorcio TLBG / UP / STRI (2003: 325, 333), Jaramillo et al. (2010: 617, 633), Köhler (2011: 130, 131), Grant et al. (2017: S 38, S 40 – S 41, S 60) Holotype. MHCH 2683 (original field number AB 026), adult male, collected by AB and Marcos Ponce on 26 July 2011 in San Lucas (8.9884, –80.5824, 150 m), Donoso District, Colón Province, Republic of Panama. Paratypes. SMF 97163 male from Donoso District, Colón Province, Republic of Panama (8.9658, –80.6250, 60 m), collected by AB and Marcos Ponce on 19 July 2011; SMF 97076 male from Chicheme, Donoso, San José del General, Omar Torrijos Herrera District, Colón Province (8.8653, –80.6438, 100 m), collected by AB on 23 January 2013; MHCH 3290 male from Cerro El Gaital, Llano Grande, La Pintada District, Coclé Province, Republic of Panama (8.7087, –80.4241, 420 m), collected by AB and Marcos Ponce on 13 November 2016. Diagnosis. Colostethus donosensis sp. nov. is characterized by the following combination of characters: (1) dorsum 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, 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 IV reaching distal subarticular tubercle of finger III, while finger I not and finger II almost; disk pads even; (7) no fringes, weak webbing between fingers I – III; (8) palmar tubercle rounded, flattened and larger than thenar tubercle; thenar tubercle low and elongated; subarticular tubercles rounded and globular, first tubercle more evident; (9) heel smooth; (10) no fringes on toes, weak webbing; (11) plantar tubercle indistinguishable, subarticular tubercles present (one on toes I and II, two on toes III and V, three on toe IV); inner metatarsal tubercle longitudinally elongated, outer metatarsal tubercle smaller; tarsal keel distinct; (12) dorsal ground color in life brown, uniform or with dark blotches, posterodorsal longitudinal dark band on thighs; dorsal ground color of limbs reddish or orangish; venter pale, white belly and pink chest, forelimbs and hind limbs (Fig. 5 B); (13) axilla-groin stripe interrupted at about a mid-distance anteriorly from groin; (14) SVL males: 18.0 ± 0.76 (17.4–18.5) mm, females: 14.4 ± 0 (14.4–14.4) mm; (15) advertisement calls at dominant frequency of 4585.7 ± 183.83 (4393–4759) Hz, with a note repetition rate of 6.0 ± 0.17 (5.8–6.1) notes / s and with a number of notes within one call being up to 33. Comparisons. Colostethus donosensis sp. nov. differs from C. panamansis in maximum SVL of adult males (18.5 mm vs. 27 mm in C. panamansis) and in degree of webbing (toes weakly vs. moderately webbed in C. panamansis). Males’ advertisement calls consist of bouts of 11–33 notes (vs. 2–5 notes per call in C. panamansis), single notes have a duration of 47 ± 4.92 ms (vs. 60 ms in C. panamansis). Colostethus donosensis 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 maximum SVL of adult males, which is 18.5 mm vs. 32.4 mm C. latinasus. Male advertisement calls differ significantly in their dominant frequency (4392.8–4558.8 Hz vs. 2658–2673 Hz in C. latinasus). From Colostethus pratti, C. donosensis sp. nov. differs by dorsal skin color, which is grey with brown speckles (unlike in C. pratti, which is uniform brown), by color pattern on the posterior surface of thighs (one longitudinal dark band on reddish background vs. pair of dark areas separated by a light line in C. pratti); throat of C. pratti males is mottled with grey to dark brown (vs. uniform greyish or yellowish in C. donosensis sp. nov.); toes weakly webbed (vs. no webbing in C. pratti); by advertisement calls emitted at a lower note repetition rate (6.0 ± 0.17 notes / s) than C. pratti (19 notes / s), and producing calls with 11–33 notes (vs. 29–129 notes per call in C. pratti). From Colostethus bocatoro sp. nov., C. donosensis sp. nov. differs by the ventral coloration, that is greyish or yellowish at throat and pink at chest and limbs (vs. uniformly reddish / pink / orange at the whole body except for white belly); by finger webbing (weak between fingers I – III vs. all fingers moderately webbed in C. bocatoro sp. nov.); advertisement calls are performed individually or with only one response to another male (vs. in a chorus with other males) and with a note repetition rate significantly lower (6.0 ± 0.17 notes / s; t = – 16.162, df = 7, p <0.001) than in C. bocatoro sp. nov. (9.5 ± 0.35 notes / s). From Colostethus kuara sp. nov., C. donosensis sp. nov. differs by the absence of yellow coloration in groins and axillae; and by calling individually or with only one response to another male (vs. in a chorus with other males). Advertisement calls have a dominant frequency of 4393–4759 Hz, 4585.7 ± 183.83 Hz, that is lower than in C. kuara sp. nov. (5140–5354 Hz, 5242.2 ± 107.03 Hz), though it is correlated with the body size. From Colostethus portobelensis sp. nov., C. donosensis sp. nov. differs by the absence of yellow coloration in groins and axillae and by the ventral coloration, that is pale with white belly and pink chest and limbs (vs. white belly, darker throat and yellowish limbs); weak webbing between fingers I – III (vs. all fingers weakly webbed in C. portobelensis sp. nov.); advertisement calls have a dominant frequency of 4393–4759 Hz, 4585.7 ± 183.83 Hz, that is higher than in C. portobelensis sp. nov. (3750–4125 Hz, 3950.9 ± 127.02 Hz), though it is correlated with the body size. Note duration is significantly lower in C. donosensis sp. nov. (47.0 ± 4.92 ms vs. 61.8 ± 5.12 ms; t = – 4.2473, df = 8, p = 0.0028) and a note repetition rate is slightly, however significantly higher (6.0 ± 0.17 notes / s; t = 5.1087, df = 8, p <0.001) than of C. portobelensis sp. nov. (5.3 ± 0.22 notes / s). From Colostethus sanblasensis sp. nov., C. donosensis sp. nov. differs by the absence of reddish or yellow coloration in groins and axillae; by a weak toe webbing (vs. only vestigial in C. sanblasensis sp. nov.); by a significantly lower note repetition rate (6.0 ± 0.17 notes / s vs. 11.7 ± 1.98 notes / s; t = 10.593, df = 7, p <0.001) and by calling individually or with only one response to another male (vs. in a chorus with other males). Colostethus donosensis sp. nov. has a significantly smaller SVL in males than C. chucantiensis sp. nov. (17.4–18.5 mm, 18.0 ± 0.76 mm vs. 18.4–25.1 mm, 21.8 ± 1.75 mm; t = 3.0199, df = 14, p <0.01); it has no yellow coloration in axilla and groin (vs. yellow coloration in groins and axillae in C. chucantiensis sp. nov.); advertisement calls have a dominant frequency of 4393–4759 Hz, 4585.7 ± 183.83 Hz, that is higher than in C. chucantiensis sp. nov. (3523–4016 Hz, 3816.5 ± 205.93 Hz), though it is correlated with the body size. Number of notes per call of 19.0 ± 12.17 (11–33) notes is higher than 5 ± 1.79 (3–8) notes of C. chucantiensis sp. nov. Measurements of 2 adult males and 1 female in the type series. SVL 18.0 ± 0.76 (17.4–18.5) mm in males, 14.4 mm in female; AGD 6.7 ± 0.68 (6.2–7.2) mm (36–39 % of SVL) in males, 6.0 mm (42 % of SVL) in female; HW 34–42 % of SVL in males, 39 % in female, and 0.8–1.1 of HL (0.9 in female); TD 38–43 % of EL in males, 37 % in female; EL 37–45 % of HL in males, 36 % in female; EL 15–16 % of SVL in males, 14 % in female; EN 9–11 % of SVL in males, 11 % in female; IO 12–13 % of SVL in males, 27 % in female; FL 23–26 % of SVL in males, 21 % in female; TL 8.2 ± 0.41 (7.9–8.5) mm (45–46 % of SVL) in males, 6.1 mm (43 % of SVL) in female. Measurements of holotype. (in mm) SVL 17.41; HL 7.35; HW 6.13; EN 1.85; IN 2.25; EL 2.74; IO 2.19; TD 1.05; FAL 4.55; TL 7.87; 3 FW 0.40; 4 FW 0.30; 3 FD 0.39; 4 FD 0.41; AGD 6.20. Description of holotype. Both dorsal and ventral surfaces smooth, except for a few folds in the sacral area. Head truncated, snout bluntly rounded in dorsal aspect and rounded to slightly protruding in lateral aspect. Nares laterally and slightly anteriorly directed, clearly visible in frontal, dorsal and ventral view. Canthus rostralis barely distinct, blunt and slightly convex. Loreal region concave. Tympanic ring is almost completely discernible externally, except for its postero-dorsal margin. Maxillary teeth present, medial lingual process absent. Relative appressed finger lengths I <II <IV <III. Fingers I and II reaching the disk of finger IV. Finger IV slightly exceeds distal subarticular tubercle of finger III. All hand tubercles well defined. Fingers I and II each with a single subarticular tubercle, fingers III and IV with two of them. Thenar tubercle elliptical, palmar tubercle rounded. Finger fringes absent. Finger III not swollen. Disks rounded, weakly expanded, with paired dorsal scutes. Relative length of toes I <II <V <III <IV. All foot tubercles well defined. Toes I and II each with a single subarticular tubercle, toes III and V with two, and toe IV with three tubercles. Webbing weak. Inner metatarsal tubercle longitudinally elongated, outer metatarsal tubercles small and rounded, tarsal keel small but visible as a dark line leading proximally from the inner metatarsal
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