New and noteworthy locality records of anurans from northeastern Andes of Colombia

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1 Herpetology Notes, volume 12: (2019) (published online on 13 January 2019) New and noteworthy locality records of anurans from northeastern Andes of Colombia Fabio Leonardo Meza-Joya 1,*, Wilfredo Chinchilla-Lemus 2, Eliana Ramos 1, Orlando Armesto 3,4, and Aldemar A. Acevedo 3,4 Tropical Andes harbour the greatest frog diversity worldwide with a high number of restricted-range species that are increasingly threatened by habitat modification and climate change (Myers et al., 2000; Grenyer et al., 2006; La Sorte and Jetz, 2010). In Colombia, the Andes Mountains are divided into three major mountain ranges: Cordillera Occidental, Cordillera Central, and Cordillera Oriental; with the later harbouring the lower anuran diversity (155 species; 21% of the national anuran diversity; Acosta-Galvis, 2017) but the higher level of endemism (77 species; ca. 50%; Lynch et al., 1997; Bernal and Lynch, 2008; Armesto and Señaris, 2017). While most anuran species from the Cordillera Oriental of Colombia have been described just over the last 50 years, mainly thanks to the taxonomic work by J.D. Lynch, P.M. Ruiz Carranza, and M.C. Ardila Robayo, the recent discovery of new species (e.g., Anganoy-Criollo, 2012; Acosta-Galvis, 2015; Ospina- Sarria et al., 2015; Rivera-Correa et al., 2016; Rojas- Runjaic et al., 2018) and the report of new localities for described ones (e.g., Duarte-Cubides and Cala-Rosas, 2012; Acevedo et al., 2014; Meza-Joya, 2016), suggest that additional surveys in poorly known areas of this cordillera and the careful examination of specimens deposited in herpetological collections are still needed to a better documentation of the regional anuran diversity. Expeditions to several localities on the northeastern portion of the Cordillera Oriental and Sierra Nevada de Santa Marta in Colombian Andes (Santander, Norte de Santander, and La Guajira departments) between August 2012 and April 2016 resulted in the finding of unreported populations of eight anuran species belonging to five families: Aromobatidae (1 species), Bufonidae (1 species), Centrolenidae (1 species), Craugastoridae (4 species), and Dendrobatidae (1 species). Specimens were identified based on the original species descriptions (Günther, ; Lynch and Duellman, 1973; Lynch, 1984, 1996; Kaplan, 1997; Barrio-Amorós et al., 2007; Anganoy-Criollo, 2012; Ospina-Sarria et al., 2015). Voucher specimens were deposited in the herpetological collection of Universidad Industrial de Santander (UIS-A) and herpetological collection of Universidad de Pamplona (MCNUP-H). We provide updated distributional maps for the reported species based on relevant literature and specimens housed at herpetological collections. Records containing uncertain or inconsistent geographic information were excluded from the maps. Collection acronyms followed Frost (2017). Family Aromobatidae 1 Colombia Endémica, Asociación para el estudio y la conservación de los recursos naturales, Bucaramanga, Colombia. 2 Grupo de Estudios en Anfibios y Reptiles de Santander (G.E.A.R.S), Bucaramanga, Colombia. 3 Programa de Doctorado en Ciencias Biológicas, Mención Ecología, Laboratorio de Biología Evolutiva, Pontificia Universidad Católica de Chile, Santiago, Chile. 4 Grupo de Investigación en Ecología y Biogeografía, Universidad de Pamplona, Pamplona, Colombia. * Corresponding author. fabio.meza@correo.uis.edu.co Allobates ignotus Anganoy-Criollo, This species was described from three localities on the western flank of Serranía de Perijá in Cesar, Colombia (Anganoy-Criollo, 2012), but recently was reported from six additional localities on this mountain range in Cesar and La Guajira departments, Colombia (Granda- Rodríguez et al., 2018). Known localities range in elevation from 400 to 1236 m (Anganoy-Criollo, 2012; Granda-Rodríguez et al., 2018). Herein we report a tenth locality based on three specimens (UIS-A ) collected at La Gran China site, vereda Barriales-Nuevas

2 62 Ideas, El Molino river, El Molino municipality, La Guajira department, Colombia ( N, W, 758 m a.s.l.). This report constitutes the northernmost locality record for this species, extending the species distribution by ca. 30 km NE from the closest locality previously reported (Nicaragua creek, La Jagua del Pilar; Fig. 1). Allobates ignotus differs from its congeners by the combination of the following characteristics (Anganoy- Criollo, 2012): (1) disks on finger III and toe IV, slightly expanded; (2) faintly differentiated lateral keels on all fingers; (3) fringes on all toes, slightly expanded; (4) pale dorsolateral stripe extended from eyes to midlevel of insertion of thigh (to the posterior-level of the of insertion of thigh in the specimens reported here), and does not drop onto thigh; (5) diffuse pale oblique lateral stripe as a series of spots (diffuse pale but not in a discrete series of spots in the specimens reported here), extending anteriorly from the inguinal region to more than middle body; (6) ventrolateral stripe present; (7) gular-chest region cream, sexually dimorphic; with scarce brown stippling in adult females and uniformly dark, brown to greyish brown, in adult males; (8) cloacal tubercles absent; and (9) pattern of coloration on dorsum brown to dark brown, with one diffuse wide band from snout to urostyle. We consider that variation in the dorsolateral and oblique lateral stripes in the specimens Figure 1. Lateral view of an alive specimen of Allobates ignotus (UIS-A 6015) and its updated geographic distribution: the black circle indicates the localities referred in Anganoy- Criollo (2012) and Granda-Rodríguez et al. (2018), and the red circle indicates the new distributional record reported here. Datum WGS84. Photo: F.L. Meza-Joya. Fabio Leonardo Meza-Joya et al. reported here (see above) is due to intraspecific variation. The revision of the photographs of four specimens from some of the localities reported in Granda-Rodríguez et al. (2018) shows evident variation in these characters, with the specimen of La Guajira department showing a colour pattern similar to the specimens reported here. Specimens were on the floor of riparian Sub-Andean forest associated to El Molino river. During fieldwork, from 13:00 to 17:00 h, we recorded one female, six males, five tadpoles, and two metamorphic specimens. Males were found calling under rocks in the margins of the river, suggesting reproductive activity. Family Bufonidae Rhaebo glaberrimus (Günther, 1869). This species is known from the eastern flank of the Cordillera Oriental in Colombia and south-eastern border of Cordillera de Mérida in Venezuela, at elevations between 300 and 1470 m (Chacón-Ortíz et al., 2001, 2002; Mueses- Cisneros et al., 2012). In Colombia, R. glaberrimus has been registered from several localities on the eastern piedmont of the Cordillera Oriental in Boyacá, Casanare, Cundinamarca, and Meta departments, between 520 and 1470 m elevation (Lynch, 2006; Mueses-Cisneros et al., 2012; but see Ruiz-Carranza et al., 1996). The type locality of this species (i.e., Bogotá, Cundinamarca ) is probably in error because Bogotá city and its surroundings are about 1100 m above the upper altitudinal limit of the species (1470 m a.s.l.), this region harbour habitats very different of those where the species occurs, and herpetological surveys in this region has not been recorded the species (Mueses-Cisneros et al., 2012). Likewise, the record from Amazonas department (Puerto Rastrojo, Mirití-Paraná river; IAvH 2502) reported by Acosta-Galvis (2017) was assigned to Rhaebo guttatus by Mueses-Cisneros et al. (2012) based on the morphological examination of the specimen. The specimens reported here (MCNUP-H 0423, 0433, 0441) comes from three sites along an elevational gradient at vereda San Antonio, San Lorenzo river, Toledo municipality, Parque Nacional Natural Tamá, Norte de Santander department, Colombia ( N, W, 646 m a.s.l.; N, W, 780 m a.s.l.). This report account for the first records of this species from Norte de Santander department and extend the species distribution by ca. 95 km SSW from the closest locality previously reported (Uribante municipality, Táchira state, Venezuela), filling the distribution gap between the previously recorded Colombian and Venezuelan localities (Fig. 2).

3 New and noteworthy locality records of anurans from Colombia 63 Figure 2. Dorsal and ventral view of a museum specimen of Rhaebo glaberrimus (MCNUP-H 0423) and its updated geographic distribution: the black circles indicates the localities referred in Chacón et al. (2001), (2002), Lynch (2006), and Mueses-Cisneros et al. (2012), and the red circles (almost grouped into a single point) indicates the new distributional records reported here. Datum WGS84. Photo: A. Acevedo. Rhaebo glaberrimus is distinguished from other congeners by the combination of the following characteristics (Günther, 1869; Chacón-Ortíz et al., 2002; Mueses-Cisneros et al., 2012): (1) preocular ridge absent; (2) cephalic crests absent; (3) enlarged parotoid glands; (4) smooth dorsal skin; (5) small skin folds on hind limbs; (6) prominent internal metatarsal tubercles on feet; (7) cloacal opening near the inferior part of the thighs in females or ventrally in males; (8) anterior part of hind limbs mottled with pale yellow; and (9) groin, axilla, and posterior part of hind limbs with pale reddish to orange spots. Individuals were recorded between 14:00 and 17:00 h, over rocks and on the base of shrubs (Miconia sp.) at the margins of San Lorenzo river. Vegetation at the study site corresponds to riparian Sub-Andean forest associated with large cattle grazing areas. Family Centrolenidae Rulyrana flavopunctata (Lynch and Duellman, 1973). This species occurs in the Amazonian-versant of the Cordillera Oriental of Colombia and Ecuador between 300 and 1850 m elevation (Lynch and Duellman, 1973; Cisneros-Heredia and McDiarmid, 2006; Lynch, 2006; Cisneros-Heredia, 2009; Almendáriz et al., 2014). In Colombia, R. flavopunctata occurs on the foothills of the Cordillera Oriental in Boyacá, Casanare, Meta, and Caquetá departments, between 540 and 1650 m elevation (Lynch, 2006; Pedroza-Banda et al., 2014; Astwood-Romero et al., 2016). Here we report this species from San Lorenzo river, vereda San Antonio, National Natural Park Tamá, Toledo municipality, Norte de Santander department, Colombia ( N, W, 714 m a.s.l; N, W, 829 m a.s.l). Collected specimens (MCNUP-H 0231, 0233) account for the first records of R. flavopunctata at foothills of Tamá Massif in Colombia and extends its distributional range ca. 224 km NNE from the closest known record (La Limonita stream, Pajarito municipality, Boyacá department; Fig. 3). Rulyrana flavopunctata can be easily distinguished from other glass frogs by the combination of the following characters (Lynch and Duellman, 1973): (1) head slightly wider than body; (2) snout short, rounded in dorsal and lateral profiles; (3) prevomerine dentigerous processes small, bearing 0-3 teeth on low processes; (4) humeral spine absent in males; (5) webbing absent between the inner fingers; (6) dermal folds on arms and legs absent; (7) three-fourths of tympanum visible; (8) dorsum green with minute pale yellow flecks; (9) edge of upper lip Figure 3. Dorsolateral view of an alive specimen of Rulyrana flavopunctata (MCNUP-H 0232) and its updated geographic distribution: the black circles indicates the localities referred in Lynch and Duellman (1973), Lynch (2006), Cisneros- Heredia (2009), Almendáriz et al. (2014), Pedroza-Banda et al. (2014), Guayasamín et al. (2015), Acosta-Galvis (2017), IAVH, QCAZ, and MUJ, and the red circles (grouped into a single point) indicates the new distributional records reported here. Datum WGS84. Photo: A. Acevedo.

4 64 pale yellow (pale green in the specimens reported here); (10) fingers and toes yellow (fingers mostly pale green in the specimens reported here); (11) parietal peritoneum white; and (12) visceral peritoneum clear. We consider that colour differences in the specimens reported here (see above) are probable due to intraspecific variation. However, further studies (morphological, acoustic, and genetic) should be conducted for this widely-distributed and highly-variable taxon as cryptic diversity may occur (Cisneros-Heredia, 2009). Some adult males were found calling from stones and perching on leaves (Cyclanthaceae) at heights until 3 m, indicating reproductive activity. Vegetation at the study site corresponds to riparian Sub-Andean forest associated with large cattle grazing areas. Family Craugastoridae Craugastor metriosistus Ospina-Sarria, Angarita- Sierra and Pedroza-Banda, This species was previously known from the middle and upper portions of the Magdalena River Valley in Colombia, with confirmed records from Antioquia, Boyacá, Caldas, Cesar, Cundinamarca, Santander, and Tolima departments, between 115 and 1150 m a.s.l (Ospina- Sarria et al., 2015; Restrepo et al., 2017). The specimens reported here (UIS-A 5911, ) comes from three sites on western flank of the Cordillera Oriental of Colombia: (i) an adult male from La Mica stream, corregimiento Otaré, Ocaña municipality, Norte de Santander department ( N, W, 1380 m a.s.l), (ii) an adult female from Cañada La Esperanza, La Esperanza neighbourhood, Bucaramanga municipality, Santander department ( N, W, 683 m a.s.l.), and (iii) a juvenile specimen from El Diviso farm, vereda La Colorada, San Vicente de Chucurí municipality ( N, W, 1280 m a.s.l). The record from Norte de Santander account for the first confirmed record of this species in this department, extending its distributional range ca. 45 km NNE from the closest known record (El Cobre farm, vereda Vega del Oso, San Martín municipality, Cesar department; Fig. 4) and its upper altitudinal limit from 1150 (Ospina- Sarria et al., 2015) to 1380 m. Craugastor metriosistus differs from all other species in the Craugastor fitzingeri group by the following combination of characters (Ospina-Sarria et al., 2015): (1) toe webbing on the outer side of toe III reaching the proximal portion of distal subarticular tubercle (III2 ⅓, III2 + or III2); (2) toe V shorter than toe III; (3) low supernumerary tubercles, restricted to the proximal Fabio Leonardo Meza-Joya et al. Figure 4. Dorsolateral view of an alive specimen of Craugastor metriosistus (UIS-A-5911) and its updated geographic distribution: the black circles indicate the localities referred in Ospina-Sarria et al. (2015) and Restrepo et al. (2017), and the red circles indicates the new distributional records reported here. Datum WGS84. Photo: W. Chinchilla-Lemus. segments of fingers; and (4) supratympanic fold distinctly curved downwards. The collected specimens slightly differ from the original description (Ospina- Sarria et al., 2015) by lacking reddish-brown colouration on the posterior surfaces of the thighs, which is herein interpreted as intraspecific variation. They also account for the largest male (Snout-vent length [SVL] = mm; UIS-A 5911) and female specimens (SVL = mm; UIS-A 6018) reported for this species (against 37.7 mm and 60.7 mm, respectively; Ospina-Sarria et al., 2015). Individuals were registered at night on the floor of secondary vegetation (corregimiento Otaré, Ocaña municipality), riparian Sub-Andean forest (La Esperanza neighbourhood; Bucaramanga municipality), and coffee plantations shaded by native trees (vereda La Colorada; San Vicente de Chucurí municipality). Pristimantis acutirostris (Lynch, 1984). This species is currently known from five localities on the northwestern slopes of the Cordillera Oriental of Colombia in Santander department, at elevations between 1740 and 2400 m (Bernal and Lynch, 2008; Frost, 2017). The locality of a specimen (ICN 5490) from Calarcá municipality, Quindío department, probably is in error (see Lynch 1984). Here we report this species from two additional localities (UIS-A 6023; PAG 948

5 New and noteworthy locality records of anurans from Colombia awaiting catalog number in the ICN): (i) El Guamo site, vereda Calichana, Mogotes municipality, Santander department, Colombia ( N, W, 1623 m a.s.l.) and (ii) Dosquebradas farm, vereda Ajizal, Moniquirá municipality, Boyacá department, Colombia ( N, W, 2150 m a.s.l.; N, W, 2230 m a.s.l.). These records are the northernmost and southernmost localities reported for this species, extending their lower elevational distribution from 1,740 (Bernal and Lynch, 2008) to 1,623 m, and accounting for its first record from Boyacá department at ca. 36 km ENE from the closest known record (vereda el Taladro, Charalá municipality, Santander department; Fig. 5). Pristimantis acutirostris can be easily distinguished from similar species by the following characteristics (Lynch, 1984): (1) dorsum shagreened, venter areolate; (2) dorsolateral folds present; (3) tympanic annulus present, small, round; (4) snout acuminate in dorsal view, rounded in profile; (5) vocal slits and subgular vocal sac present; (6) finger fringes absent; (7) small tubercle on the inner edge of tarsus; (8) toe fringes slightly, no webbing; (9) concealed surfaces of thighs yellow with brown reticulation; (10) canthal and supratympanic stripes dark brown; and (11) iris pale blue with reddish horizontal streak. Specimens were found at night in a small patch of secondary lower montane wet forest adjacent to pastures (vereda Calichana, Mogotes municipality) and in a young secondary forest with well-defined undergrowth vegetation dominated by Quercus humboldtii, next to pastures (vereda Ajizal, Moniquirá municipality). Pristimantis yukpa Barrio-Amorós, Rojas-Runjaic and Infante, This species was previously known from at least 11 localities on the eastern slope of Perijá, Zulia state, Venezuela (Barrio-Amorós et al., 2007; IUCN SSC, 2011) and one locality on western slope of the Serranía de Perijá, La Guajira department, Colombia (Meza-Joya, 2016), at elevations between 500 and 1600 m (Barrio-Amorós et al., 2007; IUCN SSC, 2011). The specimens reported here (UIS-A , 5909, 5912) comes from two sites on the western slope of Cordillera Oriental: El Lobo and La Mica farms, corregimiento Otaré, Ocaña municipality, Norte de Santander department, Colombia ( N, W, 1380 m a.s.l; N, W, 1378 m a.s.l). This report constitutes the second record of this species for Colombia and the first records from habitats outside Serranía del Perijá, extending its distributional range ca. 179 km SSW from the closest known record (Yukpa village Kiriponsa, Machiques de Perijá municipality, Zulia state, Venezuela; Fig. 6). Figure 5. Dorsolateral view of an alive specimen of Pristimantis acutirostris (UIS-A 6023) and its updated geographic distribution: the black circles indicates the localities referred in Lynch (1984), MUJ, and UIS-A, and the red circles indicates the new distributional records reported here. Datum WGS84. Photo: G. Olarte. Figure 6. Dorsolateral view of an alive specimen of Pristimantis yukpa (UIS-A-5895) and its updated geographic distribution: the black circles indicates the localities referred in Barrio-Amorós et al. (2007) and Meza-Joya (2016), and the red circles (grouped into a single point) indicates the new distributional records reported here. Datum WGS84. Photo: W. Chinchilla-Lemus.

6 66 Pristimantis yukpa can be identified by the combination of the following characteristics (Barrio-Amorós et al., 2007): (1) skin of dorsum with scattered conical tubercles; (2) skin of venter areolate; (3) upper eyelid with ill-prominent tubercles; (4) vomerine dentigerous processes small and oblique; (5) finger I slightly shorter than II; (6) discs on digits longer than wide; (7) discs on fingers III and IV larger than those on finger I and II; (8) fringes on fingers II and III; (9) two metatarsal tubercles; (10) basal webbing between toes IV and V; (11) lateral fringes on fingers II and III; (12) canthus rostralis distinct; (13) snout subacuminate in dorsal view and profile; (14) tympanum ill-conspicuous; (15) in life dorsum creamy brown withill-defined W and inverted V-shaped marks; (16) venter white immaculate; and (17) iris pale bronze with fine black reticulations. Individuals were adult males calling on leaves, stalk, and branches of coffee and fruit trees at heights between 60 and 180 cm. Tachiramantis douglasi (Lynch, 1996). This species is known from several localities on the eastern and western slopes of the Cordillera Oriental of Colombia, between 1630 and 2670 m elevation, in the departments of Santander and Norte de Santander (Lynch, 1996; Bernal and Lynch, 2008). Most of the specimens reported herein (UIS-A , MCNUP-H ) come from three paramo zones on both flanks of Cordillera Oriental in Colombia. Two sites from an elevation gradient on the western flank at paramo de Berlín, vereda Esparta, Santa Bárbara municipality, Santander department ( N, W, 3001 m a.s.l.; N, W, 3112 m a.s.l.), and two paramos on the eastern flank at Parque Nacional Natural Tamá: (i) paramo La Cabrera, vereda El Molino, Herrán municipality, Norte Santander department ( N, W, 2980 m a.s.l.) and (ii) paramo del Tamá, vereda Siberia, Herrán municipality, Norte Santander department ( N, W, 3109 m a.s.l.). These specimens account for the first records of this species for páramo habitat and extend its upper distributional range from 2670 (Bernal and Lynch, 2008) to 3112 m elevation. An additional specimen (UIS-A 6262) was collected in a patch of riparian high-andean forest at Plumajera stream, corregimiento Pangote, San Andrés municipality, Santander department ( N, W, 2486 m a.s.l). This report constitutes the southernmost locality record for this species, extending its distribution by ca. 33 km SE from the closest locality previously reported (Vereda Planadas, Piedecuesta municipality, Santander department; Fig. 7). Fabio Leonardo Meza-Joya et al. Tachiramantis douglasi can be easily distinguished from similar species by the following characteristics (Lynch, 1996): (1) short dorsolateral folds, reaching the shoulder; (2) tympanum prominent, rounded; (3) no enlarged ulnar tubercles; (4) subconical tubercle on heel, (5) inner tarsal fold present; (6) two metatarsal tubercles; (7) low supernumerary plantar tubercles; (8) large finger disks; and (9) labial stripe white. Individuals were found between 16:00 and 23:00 h perching on shrubs (Miconia sp.) and mosses (Lycopodium sp. and Sphagnum sp.), at heights from 15 to 40 cm. Adult males were calling, suggesting reproductive activity. Family Dendrobatidae Colostethus ruthveni Kaplan, This species was previously known from several localities on Sierra Nevada de Santa Marta (SNSM) in Magdalena, La Guajira, and Cesar departments, from 680 to 1500 m elevation (Kaplan, 1997; Granda-Rodríguez et al., 2008; Rueda-Solano and Castellanos-Barliza, 2010; González-Maya et al., 2011; Romero and Lynch, 2012; Blanco-Torres et al., 2014; Granda-Rodríguez et al., 2014). This species is currently considered part of the newly recognized Colostethus ruthveni group, a clade more closely related to all dendrobatines, except Figure 7. Dorsolateral view of an alive specimen of Tachiramantis douglasi (UIS-A 6020) and its updated geographic distribution: the black circles indicates the localities referred in Lynch (1996), Arroyo et al. (2003), IAVH, MHUA-A, and UIS-A, and the red circles indicates the new distributional records reported here. Datum WGS84. Photo: F. Cediel.

7 New and noteworthy locality records of anurans from Colombia 67 Phyllobates, than to any other species of the former genus Colostethus sensu lato (Grant et al., 2017). This group is composed of two species inhabiting the SNSM: Colostethus ruthveni and Colostethus sp. ruthveni-like, an undescribed species that strongly resembles C. ruthveni (Grant et al., 2017). Here we report Colostethus ruthveni from three additional localities at the eastern flank of the SNSM in La Guajira department (UIS-A ): (i) El Arroyo creek, corregimiento La Sierrita, San Juán del Cesar municipality ( N, W, 776 m a.s.l.; N, W, 941 m a.s.l.), (ii) La Vainilla site, vereda Larga La Vida, corregimiento Mingueo, Dibulla municipality ( N, W; 514 m a.s.l.), and (iii) La Concepción farm, vereda La Totumita, corregimiento Las Flores, Dibulla municipality ( N, W, 483 m a.s.l.). The report from corregimiento La Sierrita (San Juán del Cesar municipality) extend the distribution of this species ca. 28 km S from the closest known locality in La Guajira department (basins of Tapias river, south of Riohacha municipality) and account for a wide distribution of this group on the SNSM massif (Fig. 8). The locality record from corregimiento de Nabusimake, Valledupar municipality, Cesar department (ICN 35211; see Anganoy-Criollo, 2012) extend the species upper altitudinal limit from 2100 (Granda-Rodríguez et al., 2014) to 2400 m. Colostethus ruthveni differs from other members of this genus by the combination of the following characters (Kaplan, 1997): (1) narrow dorsolateral pale stripe present; (2) upper part of flank dark stripe-like; (3) oblique lateral stripe, throat collar, chest spots, absent; (4) basal webbing on feet, absent on hands; and (5) expanded finger and toe discs. It is important to note that specimens reported here are tentatively treated here as Colostethus ruthveni because they fit entirely with the description of the species provided by Kaplan (1997). However, further morphological, acoustic, and genetic studies are required to clarify the taxonomic status of species in the Colostethus ruthveni group. During fieldwork we recorded 23 individuals, including adult female, adult males, and metamorphs. All specimens were found on rocks on the floor of riparian Sub-Andean forests. Adult males were calling, indicating reproductive activity. Discussion Our findings confirm that anuran diversity in northeastern Andes of Colombia is still underestimated Figure 8. Dorsolateral view of an alive specimen of Colostethus ruthveni (UIS-A 5479) and its updated geographic distribution: the black circles indicates the localities referred in Kaplan (1997), Granda-Rodríguez et al. (2008), Rueda-Solano and Castellanos-Barliza (2010), González-Maya et al. (2011), Anganoy-Criollo (2012), Romero and Lynch (2012), Blanco-Torres et al. (2014) and Granda-Rodríguez et al. (2014), the red circles indicates the new distributional records reported here, and the blue circle indicate the locality for Colostethus sp. ruthveni-like reported by Grant et al. (2017). Datum WGS84. Photo: F.L. Meza-Joya. and poorly known. This region is located near to the international border with Venezuela, sharing with this country a series of massifs and mountain ranges that represents an ecological and geological continuum (e.g., Serranía or Sierra de Perijá and Tamá Massif), thus, anuran species previously thought to be restricted to Venezuelan Andes are now known to occur in Colombian Andes (e.g., Acevedo et al., 2014; Meza- Joya, 2016) and vice versa (e.g., Rojas-Runjaic et al., 2012). Records from the species reported here offer a more detailed picture of its distributional range along altitudinal and latitudinal gradients. Further herpetological surveys and detailed revision of museum specimens certainly will lead to new records and to the discovery of undescribed species and cryptic diversity. Acknowledgments. Field work was funded by Corpoguajira, Conservación Internacional Colombia, Asociación Selva, Asociación Colombia Endémica, Parque Nacional Natural Tamá, and Conservation Leadership Programme (Project ID , Project ID , Project ID ). Collection of Specimens was authorized by the Corporación Autónoma Regional de La Guajira (Corpoguajira, Acuerdo 0021-

8 ), Corporación para la Defensa de la Meseta de Bucaramanga (CDMB, Resolución ), Autoridad Nacional de Licencias Ambientales (ANLA, Resolución ), and Corporación Autónoma Regional de la Frontera Nororiental (Corponor, Resolución ). We are grateful to M. Rada for allowing the use of their unpublished locality record for Pristimantis acutirostris from Boyacá department and to A. Acosta who kindly provided us some locality records for Rulyrana flavopunctata. We thank M. Anganoy, M. Rada, M. Rivera, J. Ospina, J. Lynch, and S. Arroyo for corroborated species taxonomic identification and C. Hernández, F. Cediel, R. Franco, K. Silva, L. Peña, D. Gutiérrez, S. Alvarez, and A. Gallardo for field assistance. We also thank G. Olarte and F. Cediel for allowing the use of their photographs. For comments on this manuscript, we thank A. Acosta and two anonymous reviewers. References Acevedo, A.A., Pallares, R.F., Pérez, K.S. (2014). 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