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Nymphargus pijao sp. nov. (Figs. 8, 9, 10A–B, 11B, 12A–B, 13, 14, and 15) Centrolenella griffithsi Lynch & Duellman, 1973: 33; Duellman & Burrowes, 1989: 10 Cochranella griffithsi Ruiz-Carranza & Lynch, 1991a: 97 Nymphargus griffithsi Cisneros-Heredia & McDiarmid, 2007: 34 Nymphargus sp. Guayasamin, Cisneros-Heredia, McDiarmid, Peña & Hutter, 2019: 6 Common name. English: Pijaos’ glassfrog, Spanish: Rana de cristal de los Pijaos. Generic allocation. Based on our phylogenetic results, we include the new species in the genus Nymphargus. The following morphological characters further support its placement within the genus: absence of iridophores in the hepatic and intestinal peritonea, absence or reduced webbing between fingers III and IV, and absence of humeral spines in adult males (Cisneros-Heredia & McDiarmid 2007; Guayasamin et al. 2009). Holotype: Adult male, ARUQ 1377 (Figs. 8A and 9), collected in the stream Sopladero, at 40 km northwest of Corregimiento Uribe, cabaña La Romelia, Parque Nacional Natural Munchique, municipality of El Tambo, department of Cauca, Colombia (2°56′2″N, 76°56′19″W, 2267 m asl), on 17 September 2017 by Gustavo González-Durán. Paratypes: Adult males (ICN 26013–26015, 26018–23, 26025–30, 26067), and adult female (ICN 26017), collected in the stream Sopladero, at 40 km northwest of Corregimiento Uribe, Parque Nacional Natural Munchique, municipality of El Tambo, department of Cauca, Colombia (2°56′2″N, 76°56′19″W, 1890 m asl), on 01 October 1990 by Pedro M. Ruiz-Carranza. Adult males (ARUQ 1378–1379) collected at Sector La Romelia, Parque Nacional Natural Munchique, municipality of El Tambo, department of Cauca, Colombia (2°42′26″N, 76°52′58″W, 1721–2267 m asl) on 17 September 2017 by Gustavo González-Durán. Adult males (ICN 16080, 16081) and adult female (ICN 16007) collected at road from Corregimiento Nutibara to Corregimiento La Blanquita, municipality of Frontino, department of Antioquia, Colombia (6°47′55″N, 76°15′3″W, 2000 m asl) by Pedro M. Ruiz-Carranza. Adult male (ICN 16022) collected at road from Corregimiento Nutibara to Corregimiento La Blanquita, municipality of Frontino, department of Antioquia, Colombia (6°45′35″N, 76°18′22″W, 1960 m asl), on 12 July 1987 by Pedro M. Ruiz-Carranza. Adult male (MPUJ ANFB 10968) collected at municipality of Ciudad Bolivar, department of Antioquia, Colombia (1859 m asl), in February 2008 by Taran Grant, Juan Camilo Arredondo and Marco Rada. Referred material. Adult male (ICN 59946) and adult female (ICN 59949) collected at stream La Vinagra, Alto Tominejo (on trial to Corregimiento Carauta), vereda Curadiente-Chachafrutal, Corregimiento Nutibara, municipality of Frontino, department of Antioquia, Colombia (6°43′48.2″N, 76°16′2.2″W, 2050 m asl), on 26 and 27 August 2014 by Marco Rada. Adult male (ICN 16057) collected at road from Corregimiento Nutibara to Corregimiento La Blanquita, municipality of Frontino, department of Antioquia, Colombia (6°46′54″N, 76°15′57″W, 1430 m asl), on 18 July 1987 by Pedro M. Ruiz-Carranza. Adult males (ICN 16075, 16085) collected at road from Corregimiento Nutibara to Corregimiento La Blanquita, municipality of Frontino, department of Antioquia, Colombia (6°47′55″N, 76°15′3″W, 2000 m asl) by Pedro M. Ruiz-Carranza. Adult male (ICN 19730) collected at stream Las Canoas, Parque Nacional Natural Las Orquideas, vereda Calles, municipality of Urrao, department of Antioquia, Colombia (6°30′44″N, 76°14′7″W, 1770–1800 m asl) on 25 May 1988 by Pedro M. Ruiz-Carranza. Adult males (ICN 19728– 29) collected at stream Las Canoas, Parque Nacional Natural Las Orquídeas, vereda Calles, municipality of Urrao, department of Antioquia, Colombia (6°30′44″N, 76°14′7″W, 1770–1800 m asl), on 25 May 1988 by Pedro M. Ruiz-Carranza. Adult male (ICN 19727), and adult female (ICN 19635) collected at stream El Silencio, Parque Nacional Natural Las Orquideas, vereda Calles, municipality of Urrao, department of Antioquia, Colombia (6°30′46″N, 76°16′24″W, 1480–1540 m asl), on 23 May 1988 by Pedro M. Ruiz-Carranza. Adult males (ICN 24933, 24937– 38) collected at Bremen Reserve, vereda El Roble, municipality of Filandia, department of Quindío, Colombia (4°40′2″N, 75°37′42″W, 1930 m asl), on 07 November 1990 by Pedro M. Ruiz-Carranza. Adult male (ICN 9922) and adult female (ICN 9921), collected near at hacienda San Julian and hacienda El Brillante, vereda San Julián, municipality of Calarcá, department of Quindío, Colombia (2030–2100 m asl), on 31 May 1981 by John D. Lynch. Adul male (ARUQ 491), collected at farm El Jardin, vereda San Rafael municipality of Calarcá, department of Quindío, Colombia (4º30′23.81′′N, 75º36′30.081′′W 1915 m asl), on 23 July 2015 by Carlos Londoño-Guarnizo. Adult females (ARUQ 568–569), collected at farm El Jardin, vereda San Rafael municipality of Calarcá, department of Quindío, Colombia (4º30′23.81′′N, 75º36′30.081′′W 1915 m asl), on 26 September 2015 by Carlos Londoño-Guarnizo. Adult male (ARUQ 743), collected at reserve Las Cascadas de Río Verde, municipality of Cordoba, department of Quindío, Colombia (4º24′048′N, 75º39.271′W, 2096 m asl), on 05 February 2017 by Fernando Vargas-Salinas. Adult males (ARUQ 647, 648) collected at creek El Impes, Bellavista reserve, municipality of Pijao, department of Quindío, Colombia (4º19′31.8′′N, 75º42′0.36′′W, 1926 m asl), on 06 August 2016 by Fernando Vargas-Salinas. Adult males (ARUQ 888) collected at creek El Impes, Bellavista reserve, municipality of Pijao, department of Quindío, Colombia (4º19′31.8′′N, 75º42′0.36′′W, 1926 m asl), on 02 June 2018 by Fernando Vargas-Salinas. Adult males (ARUQ 1360) collected at creek El Impes, Bellavista reserve, municipality of Pijao, department of Quindío, Colombia (4º19′31.8′′N, 75º42′0.36′′W, 1926 m asl), on 15 February 2021 by Fernando Vargas-Salinas. Tadpoles (ARUQ 1235–1245, MPUJ ANFB 10967) collected at creek El Impes, reserve Bellavista, municipality of Pijao, department of Quindío, Colombia (4º19′31.8′′N, 75º42′0.36′′W, 1926 m asl), between June 2018 and January 2019 by Fernando Vargas-Salinas. Adult males (ICN 30139, 30145) collected at road near to Corregimiento of San Antonio del Chamí, municipality of Mistrató, department of Risaralda, Colombia (5°25′12″N, 75°56′18″W, 1550 m asl), on 06 April 1992 by Pedro M. Ruiz-Carranza. Adult female (ICN 30818) collected at stream Santa Helena, vereda La Trinidad, municipality of Pueblo Rico, department of Risaralda, Colombia (1540 m asl), on 01 September 1991 by Maria Cristina Ardila. Adult female (ICN 31313) collected at stream Santa Helena, vereda Los Planes, municipality of Santuario, department of Risaralda, Colombia (2430 m asl), on 01 January 1992 by Maria Cristina Ardila. Adult male (ICN 31323) collected at stream Risaralda, vereda La Cumbre, municipality of Apia, department of Risaralda, Colombia (5°8′50″N, 76°0′33″W, 2150 m asl), on 01 January 1992 by Maria Cristina Ardila. Adult male (ICN 28787) collected at Corregimiento El Boquerón, municipality of El Cairo, department of Valle del Cauca, Colombia (1900 m asl), on 22 June 1991 by John D. Lynch. Adult male (ICN 41394) collected at stream La Seca, 8 km near Corregimiento Queremal, municipality of Dagua, department of Valle del Cauca, Colombia (1940–2050 m asl), on 13 July 1995 by John D. Lynch (also see Appendix V for species and specimens examined). Characterization. Nymphargus pijao is diagnosed by the following characteristics: (1) vomerine teeth absent or present (one or two vomerine teeth) on very reduced dentigerous processes of the vomer; (2) truncate snout in dorsal and lateral view (Figs. 9A and 10A–B); (3) tympanum visible (tympanic membrane and annulus differentiated) with supratympanic fold present; (4) dorsal skin in reproductive males smooth with abundant microspicules, low warts associate with yellow spots (in life); microspicules absent in females (Fig. 8A–D); (5) ventral skin and thighs granular and translucent, pair of large, round, flat tubercles on ventral surfaces of the thighs; (6) parietal peritoneum covered by iridophores, more than 50% (condition P3, sensu Cisneros-Heredia & McDiarmid 2007); visceral peritoneum translucent with the exception of the pericardium that is covered by iridophores (condition V 1, sensu Cisneros-Heredia & McDiarmid 2007) (Fig. 8B–C); (7) liver tetra-lobed (Condition H0, sensu Cisneros-Heredia & McDiarmid 2007); (8) humeral spine absent (crista ventralis very low; Fig. 11B); (9) webbing between fingers I–II absent; basal between fingers II– IV: II (2 - – 31/2)–(3 + – 31/2) III (2 - –3 +)–(1 + –3 -) IV (Fig. 12A); (10) webbing between toes I (1 + –2 +)–(2 - – 21/2) II (1 - – 21/2)–(1 + – 21/2) III (1 - –2 +)–(1 + – 31/2) IV (11/4 – 31/2)–(1 + –2) V (Fig. 12B); (11) ulnar folds present, translucent and low; (12) nuptial excrescence type I, concealed prepollex; (13) Finger I shorter than Finger II ( Finger I length 70–95% of Finger II); (14) diameter of eye larger than width of disc on Finger III (disc of Finger II corresponds to a 31.5–61.8% of the diameter of the eye); (15) in life, dorsum green with tiny clusters of melanophores forming small flecks and yellow or green-yellowish dots; green bones (Figs. 8A and 13D); (16) color in preservative, dorsum lavender or pale lavender with or without clusters of melanophores forming small flecks and cream-colored dots (Figs. 9 and 10A–B); (17) iris coloration in life, silver white or faint gold with fine reticulations and small flecks on the upper and lower side of the iris; (18) melanophores present and abundant at the base and along fingers III and IV, less dense on Finger II, and rarely present on Finger I, present and abundant along toes IV and V, less dense in the other fingers; (19) males call from the upper surfaces of leaves (Fig. 13D) (Arcila-Pérez et al. 2017); (20) the advertisement call is one pulsed note with a duration of 0.023 ± 0.007 s (0.013 – 0.037 s, n = 7) and a dominant frequency of 3.68 ± 0.09 kHz (3.62–3.79 kHz, n = 7) (Arcila-Pérez et al. 2017); (21) unknown combat behavior, (22) egg masses deposited on the underside of leaves, although one egg clutch was found on the upper side of a leaf (Arcila-Pérez et al. 2017); clutch size: 35– 50 eggs (42 ± 6.6, n = 11) (Fig. 13A–B); (23) tadpole snout rounded in lateral and dorsal view; spiracle sinistral, located posterolaterally; tail long, low; dorsal fin slightly higher than ventral fin; mouth anteroventral; labial tooth row formula 2(2)/3(1) and occasionally 2(2)/3(1)(2) or 2(2)/2(1); upper jaw sheath arch-shaped; lower jaw sheath U-shaped; tadpoles are almost translucent, coiled intestines visible through the skin, liver orange-pink, heart, sinus hyobranchialis and other parts of the circulatory system pink (Figs. 14 and 15); (24) SVL in adult males 24.6 ± 1.4 (22.1–27.3 mm, n = 27), and in adult females 24.7 ± 1.5 (24.6–28.6 mm, n = 7) (Table 1); (25) insertion of musculus pectoralis abdominalis deep (see pectoral and humeral musculature section); (26) insertion of musculus deltoideus episternalis ventrolateral (see pectoral and humeral musculature section) (Fig. 3E); (27) small testis size (relative length with respect to that of kidneys in reproductive adults) (Fig. 4A). Description of the holotype. An adult male in good state of preservation (Fig. 9). Body size moderate (SVL = 22.9 mm); head wider than the body; HW 34.5% of SLV; head wider than long, HW/HL = 1.0; snout truncate in dorsal profile and lateral profile; END 73.1% of ED; loreal region concave, nostrils elevated producing a slight depression in the internarial area; canthus rostralis round; ED 36.7% of HW; supratympanic fold distinct, tympanic annulus distinguished and round; tympanic membrane differentiated and partially pigmented, IN 48.9% of ED; vomerine teeth absent; tongue round, slightly longer than wide, with posterior distal margin slightly notched, posterior edge not adherent to the floor of the mouth. Humeral spine absent; ulnar tubercles absent; ulnar fold present, not enameled, low; Finger I shorter than Finger II, discs round to subtruncate and wider than tympanum, the diameter of the disc of Finger III equivalent to 53.1% of ED; concealed prepollex. Hand webbing formula I-II (absent), II-III (absent), III 3 - – 21/4 IV; subarticular tubercles round, supernumerary tubercles at base of fingers, palmar tubercle slightly longer than wide, thenar tubercle elliptical, longer than palmar tubercle. Hind limbs slender, almost the same width of forearm, TL 61.8% of SLV; tarsal tubercles absent; translucent tarsal fold present (not enameled), low; inner metatarsal tubercle elliptic, outer metatarsal tubercle absent; subarticular tubercles round; webbing formulae on feet I 2 - – 21/3 II 11/4 – 21/4 III 2 + –2 + IV 21/4 –2 - V; toe discs narrower than finger discs. Smooth dorsal texture with seven low warts associated with pale light spots (in life, white in preservative) and with clusters of melanophores forming small flecks, dorsal skin smooth with abundant microspicules (only visible under stereo microscope), low warts associate with yellow spots in life, and white in preservative; ventral skin on belly and thighs granular; cloacal opening directed posteriorly, concealed by a dermal fold; pair of enlarged flat warts on the posteroventral surface of thighs. Measurements of holotype (in mm). SVL = 22.9; Head width = 7.9; Head length = 7.6; Interorbital distance = 2.8; Eye-nostril distance = 2.1; Internarial distance = 1.4; Width disc of Finger III = 1.8; Tibia length = 14.1; Foot length = 10.9; Eye diameter = 2.9; Tympanum diameter = 1.1; Hand length = 8.9; Forearm length = 5.7. Coloration of the holotype. Dorsum light green in life, with tiny clusters of melanophores forming small flecks and some yellow or cream-yellowish dots, concentrated on the dorsal surface of the head and body. Parietal peritoneum covered by iridophores, more than 50%. Visceral peritoneum translucent except for the pericardium that is covered by iridophores (M. Rada pers. obs, on freshly collected type series). Slightly pale-yellow toes and hands with few melanophores on dorsal surfaces. The iris is silver with fine reticulations and small punctuations. The bone color is green (Fig. 8A). In ethanol, dorsal surfaces of head and dorsum are pale lavender-colored with large light cream-colored dots, with dark lavender clusters of melanophores forming small flecks and cream-colored spots. Melanophores are present on fingers III and IV, less dense on Finger II, and abundant along toes IV and V, less dense in the other fingers. Translucent testis. Dark lavender iris with darker reticulations and small flecks (Figs. 9 and 10A–B). Variation. Adult individuals vary from dark green to light green with or without clusters of melanophores forming small flecks. Orange-yellowish or light-yellow dots are present on the dorsal surfaces of the head and body. Yellowish to pale green toes and hands with melanophores on dorsal surfaces. The iris ranges in color from silverwhite to faint gold with fine reticulations and minute rounded flecks. Dorsal color of specimens in ethanol varies from lavender to pale lavender-colored with different concentrations of minute dark lavender flecks and light cream-colored dots. Melanophores present on fingers III and IV, are less dense on finger II, and abundant along toes IV and V, less dense in the other fingers (Fig. 8A, E–G). Measurement and body proportion are shown in Table 1. Variation in webbing of hands and feet are shown in Table 2. Vomerine odontophores vary from absent to present. Bioacoustics. According to Arcila-Pérez et al. (2017), the advertisement call of N. pijao is a pulsed note with one or more amplitude peaks, the call duration is on average 0.023 ± 0.007s (range = 0.013 –0.037 s) with a dominant frequency of 3.68 ± 0.09 kHz (3.62–3.79 kHz). In-situ tadpole rearing. We collected a clutch with 49 embryos at the developmental Stage 9–11 sensu Gosner (1960) and Salazar-Nicholls & del Pino (2015). The in-situ management and care provided to tadpoles of N. pijao was successful and it was possible to record individuals on distinct stages. The time between the beginning of rearing tadpoles (i.e., egg hatching) and metamorphosis (Fig. 13C) was approximately 10 months (from March 2019 to January 2020). Comparative diagnosis. Nymphargus pijao can be distinguished from other centrolenids by its truncated snout (both in dorsal and lateral profile; Fig. 10); medium body size of adults (SVL in males = 22.1–27.3 mm, n = 27, in females = 24.6–28.6 mm, n = 7; Table 1); humeral spine absent (crista ventralis small in reproductive males); vomerine teeth absent or present (1–2) on very reduced dentigerous processes of the vomer; dorsum green with tiny clusters of melanophores and yellow or green-yellowish spots; skin texture smooth with minute spicules (only visible under magnification); webbing between the outer fingers absent, webbing of foot reduced (Fig. 12), and parietal peritoneum white in more than 1/2, visceral peritoneum translucent (not covered by iridophores). Among the species of Nymphargus (Fig. 16), N. pijao is most similar to N. buenaventura, N. cristinae (Ruiz-Carranza & Lynch, 1995b), N. griffithsi, and N. lasgralarias. However, N. pijao is distinguishable from these species because the snout shape in N. cristinae is subacuminated in dorsal profile and adult males are slightly larger (although their overlap in size; SVL in adult males 26.0– 31.1 mm; n = 12); N. buenaventura has iridophores covering renal capsules (white kidneys = iridophores present) and is uniformly green with small scattered pale-yellow spots (see Fig. 16 dorsum green with tiny clusters of melanophores and yellow or green-yellowish spots in N. pijao); N. lasgralarias has a dorsum uniformly green lacking spots (Guayasamin et al. 2020). Finally, reproductive males of N. griffithsi have moderate to enlarged crista ventralis of the humerus (here conside |
