Amphibians and reptiles from two localities in the northern Andes of Colombia

The northern Andes are a geographically, climatically, and ecologically complex area that harbors a high number of species and presents a high level of endemism. However, the landscape of this region has been highly transformed by human activities, including agriculture, cattle grazing, urbanization, and recently, the construction of large infrastructure projects. This study presents a list of amphibians and reptiles compiled from field surveys carried out between 2006 and 2016 in the protected areas surrounding 2 hydroelectric projects located on the eastern flank of the northern Cordillera Central in Colombia. The checklist is comprised of 101 species (43 amphibians and 58 reptiles), including 17 endemic to Colombia and 4 threatened species. The high species density, high representativeness of the regional biodiversity, and high level of amphibian endemism make these 2 protected areas important for conservation of the regional herpetofauna.


Introduction
The Andes mountain range extends for 5000 km along the western coast of South America and 3 main domains are recognized: southern, central and northern Andes (Gregory-Wodzicki 2000).The northern Andes reach their maximum geomorphological complexity in Colombia, where 3 mountain chains (the Western, Central, and Eastern cordilleras), are separated by the sedimentary basins of the Cauca and Magdalena rivers (Kattan et al. 2004).Three major historical events have shaped the biological diversity in this region: (1) the uplift of mountain ranges in a complex series of orogenic processes generating new environments and causing vicariant events, (2) the final closure of the Isthmus of Panama that allowed the biotic interchange between 2 continents, and (3) the climatic fluctuations during the Pleistocene that lead to cycles of range contraction and expansion resulting in the fragmentation and isolation of natural populations, with subsequent speciation and radiation events (Duellman 1999, Kattan et al. 2004).Thus, the diversity of ecological settings generated by this physiographic complexity has made the northern Andes one of the most biodiverse regions on the planet with an exceptional number of endemic plants and vertebrates (Fjeldsa 1994, Duellman 1999, Myers et al. 2000).
The northern Andes are also under continuous and intense anthropogenic pressure.About 70% of the human population in Colombia is concentrated in this region and agricultural activities, cattle grazing and infrastructure development (i.e.urbanization, dam construction, and roads) are among the main causes of habitat loss and fragmentation (Etter andvan Wyngaarden 2000, Fjeldsa et al. 2005).On the eastern flank of the northern Cordillera Central, several hydroelectric projects involving dam constructions, roads, and power lines were developed in the last 2 decades.As a mitigation strategy, Colombia's environmental legislation demands that these large infrastructure projects compensate the negative impacts through the implementation of effective conservation and restoration plans by preserving and monitoring the biodiversity in the surrounding areas (Minambiente 2012).As a consequence, these habitats, as virtually the only remnants left, represent a last-minute opportunity to make complete species checklists for the northern portion of the central Andes of Colombia.
Based on more than a decade of field surveys, we provide the species list of amphibians and reptiles found in the protected areas of 2 main hydroelectric projects located on the northeastern Cordillera Central in Colombia.

Methods
Study sites.This study was conducted in the northern Andes at 2 sites above 500 m elevation in the Cordillera Central in Antioquia department, Colombia (Fig. 1).The Jaguas site (06°26ʹ06.0ʺN, 075°00ʹ57.6ʺW) includes a 4000 ha private protected area with habitats in different successional stages, within the premontane wet forest life zone (bmh-PM, Holdridge 1964).The altitude ranges between 1100 and 1300 m above sea level.The temperature ranges between 18 and 24 °C and the annual rainfall between 2000 and 4000 mm.The San Carlos site (06°12ʹ57.6ʺN, 074°51ʹ21.6ʺW) includes a private protected area of 3000 ha, within the tropical moist forest life zone (bh-T, Holdridge 1964).The altitude is between 600 and 950 m above sea level.It has a mean temperature of 24 °C and the annual rainfall ranges between 2000 and 4000 mm.The 2 sites are under a bimodal rainfall regime with 1 rainy season between March and May and 1 between September and November (Poveda et al. 2005).

Data collection.
Between 2006 and 2016, the complete species inventory has been an ongoing goal as part of the monitoring plan implemented by the company ISAGEN S.A. Amphibians and reptiles were searched for in all possible microhabitats including diurnal and nocturnal sampling events during rainy, dry, and transition seasons.In total, 49 sampling points were distributed in different vegetation types.
Collected specimens were euthanized with 2% Roxicain ® , fixed in 10% formalin solution, and preserved in 70% ethanol following the protocol by Cortez et al. (2006).Voucher specimens are deposited in the Museo de Herpetología Universidad de Antioquia (MHUA), in Medellín, Colombia (Instituto Alexander von Humboldt registry # 80, since the year 2000).All procedures were conducted under permit No. 112-0046 granted by the Corporación Autónoma Regional CORNARE.We list the collected and examined specimens in Table 1 (amphibians) and 2 (reptiles).All coordinates use the WGS84 datum.
Amphibian nomenclature follows Frost (2016) and reptile nomenclature follows Uetz and Hošek (2016).We reviewed the original descriptions of the species along with the MHUA reference collection to corroborate the taxonomic identifications.We included in the list only the species having voucher specimens or photographs that allow unambiguous identification of taxa.We follow the species distribution provided by IUCN (2016) for amphibians and Uetz and Hošek (2016) for reptiles.

Amphibians
From the 2 protected areas, we report 43 amphibian species representing 13 families and 25 genera (Table 3).In the protected area of the Jaguas locality, we report 37 amphibian species from 11 families.The most diverse lineages were Hylidae with 12 species (32%) and Craugastoridae with 6 species (16%).Eleven species are endemic to Colombia (30%), and according to the IUCN Red Lists, 1 is classified as Endangered, 2 as Vulnerable, and 1 as Near Threatened (IUCN 2016).In comparison, the San Carlos site has 34 amphibian species grouped in 11 families.Again, the most diverse groups were Hylidae with 11 species (32%) and Craugastoridae with 6 species (18%).Eleven species from San Carlos locality are endemic to Colombia (32%) and 2 are listed as Vulnerable (IUCN 2016).Jaguas and San Carlos localities shared 28 amphibian species.(Rivero & Serna, 2000): Figure 2A  Material examined.Table 1.
Small glass frog which exhibits complete ventral transparency; transparent pericardium; white hepatic and visceral peritonea; white bones; iris silvery at the center, yellow at the edges with numerous black spots (Guayasamin et al. 2009).Hyalinobatrachium aureoguttatum is superficially similar to H. valeroi.Nevertheless, in preservative H. aureoguttatum differs from H. valeroi in having usually either well-defined rounded spots or elongate blotches of bright white on yellow background which correspond to clumps of iridophores (absent in H. valeroi).

Material examined. Table 1.
A medium-sized frog characterized by having extensive toe webbing and a uniformly reddish brown coloration on the posterior surfaces of the thighs.Craugastor metriosistus differ from C. longirostris and C. raniformis (the latter also distributed in the Magdalena river valley) in having the throat with median white gular stripe and dark coloration on either side (throat with spotted pattern or short lines in C. longirostris and white to cream in C. raniformis), in having posterior surfaces of the thighs reddish brown (dark to reddish brown with small and conspicuous pale spots in C. raniformis), and in having webbing extending to the proximal portion of the distal subarticular tubercle on the outer side of toe III (webbing extends to middle portion between basal and distal subarticular tubercle on the outer side of toe III in C. raniformis and C. longirostris; Ospina-Sarria et al.

2015).
Pristimantis fallax (Lynch & Rueda-Almonacid, 1999): Figure 3D Eleutherodactylus fallax Lynch and Rueda-Almonacid 1999: 308.Pristimantis fallax - Hedges et al. 2008: 118.Material examined.Table 1.Pristimantis fallax is a moderate size frog with a shagreen dorsum; finger I longer than finger II; toe V slightly longer than toe III; absence of toe webbing and broad disks on outer fingers.The most distinctive feature of P. fallax is the coloration pattern of the throat consisting in a lateral portion reticulated with gray to black, defining cream central raphe, a feature shared with Craugastor fitzingeri.Although superficially resembling C. fitzingeri, P. fallax is easily distinguished in having toe V slightly longer than toe III (toe III slightly longer than toe V in C. fitzingeri), absence of webbing on toes (basally webbed in C. fitzingeri), and by having the posterior surfaces of the thighs unicolor (with pale spots in C. fitzingeri).

Material examined. Table 1.
Pristimantis gaigei is a small to medium-sized frog characterized by having a distinct tympanum; annulus partially concealed dorsally and posteriorly by supratympanic fold; skin of dorsal surfaces shagreened; skin of venter smooth or feebly granular along lateral and posterior borders of ventral disk; discoidal folds and prominent arm slender (Lynch 1980).This species is easily distinguished from related species (i.e., P. penelopus) also distributed in the Magdalena river valley by having smooth and dark gray ventral surface, a bright yellow band on the forearm, and no calcar tubercle (in contrast with brown ventral surface with cream flecking, absence of yellow band and presence of calcar tubercle in P. penelopus).

Material examined. Table 1.
A moderated-sized frog characterized by a wide variation in color; absence of nuptial pads, discoidal fold and conical calcar tubercles; flanks and belly white to cream without blotches as well as iris yellow ocher to copper with thick brown reticulation and cream sclera (Rivera et al. 2014).This species co-occurs in the Magdalena river valley with Pristimantis penelopus and P. viejas and may be distinguished from both by having smooth skin and lacking a calcar tubercle (dorsum with tubercles including the heel in P. penelopus and P. viejas) and the concealed surfaces of the thighs yellow without blotches (orange spots on the concealed surfaces of the limbs in P. viejas; yellow spots on the concealed surfaces of the limbs, sometimes forming a reticle, in P. penelopus).

Material examined. Table 1.
A moderated-sized frog with a shagreen dorsum; tubercles posteriorly and on flanks; no dorsolateral folds; skin of venter brown with cream flecking; discoidal folds prominent; subanal tubercles only slightly larger than granules on underside of thighs; and iris copper with black reticulation.Males have spots on concealed surfaces of the thighs, a feature shared with Pristimantis viejas; nevertheless, these spots are usually yellow (sometimes forming a reticle) in P. penelopus and orange in P. viejas.Pristimantis penelopus is also similar to P. latidiscus from which it may distinguished in having a brown venter with cream flecking (in contrast to a cream venter with brown reticulation) (Lynch andRueda-Almonacid 1999, Restrepo et al. 2017).Pristimantis taeniatus (Boulenger, 1912): Figure 3H Eleutherodactylus taeniatus Boulenger 1912: 188.Pristimantis taeniatus -Hedges et al. 2008: 121.Material examined.Table 1.(Lynch & Rueda-Almonacid, 1999): Figure 4A Eleutherodactylus viejas Lynch andRueda-Almonacid 1999: 311. Pristimantis viejas -Hedges et al. 2008: 121.Material examined.Table 1.A small-sized frog with numerous non-conical tubercles on dorsum; absence of dorsolateral folds; finger I shorter than finger II; toe V very long; brown above with darker brown markings; venter cream stippled with brown and posterior surfaces of the thighs brown with large cream spots.Superficially similar to Pristimantis taeniatus in lacking a canthal stripe, but readily distinguished from that species in having orange spots on the concealed surfaces of the limbs (absent in P. taeniatus; Lynch and Rueda-Almonacid 1999).(Cope, 1868): Figure 4B Prostherapis inguinalis Cope 1868: 10.Colostethus inguinalis -Savage 1968: 745.

Material examined. Table 1.
A moderate-sized dendrobatid with a pale oblique lateral stripe extending from groin only to around midbody; pale ventrolateral stripes present; pale dorsolateral stripes absent and toes moderately webbed.Colostethus inguinalis may be distinguished from C. aff.fraterdanieli (also distributed in the Magdalena river valley) by having an incomplete pale oblique lateral stripe (complete in C. aff.fraterdanieli) and by the throat coloration of adults (black in adult males, white unpigmented or faintly pigmented gray or brown in adult females in C. inguinalis; spotted in adult males and immaculate in adult females in C. aff.fraterdanieli).It differs from C. pratti both in toe webbing (absent in C. pratti) and adult male throat coloration (pale gray or brown with irregular white spots, forming a faint, mottled or reticulated pattern in C. pratti).

Material examined. Table 1.
Small frog with skin of dorsum smooth; tympanum round and prominent; snout subacuminate in dorsal view, rounded in lateral view; all fingers and toes with expanded disks; pads triangular on toes III-IV; dorsum brown with darker blotches and venter dark brown with whitish flecks.Diasporus anthrax differs from other species of the genus by its coloration which consist in a dark venter with white flecks and pale blotches (red in life) on the concealed surfaces of the limb (orange above, venter immaculate, and lacking distinctive coloration marks on limbs in D. gularis; brown above, venter white with brown marbling, and lacking distinctive coloration marks on limbs in D. tinker) and the rounded disk pads and disk covers except for finger III and toes III-IV in contrast with all fingers bearing narrow disks and disk cover of finger III bearing elongate papilla in D. tinker; unornamented disk covers and knobbed disk cover occur in about equal frequency in D. gularis (Lynch 2001).(Boulenger, 1898): Figure 4F Hylodes gularis Boulenger 1898: 121.Diasporus gularis - Hedges et al. 2008: 47.

Material examined. Table 1.
A small frog with short body; snout nearly truncate at tip when viewed from above, truncate in profile; nostrils strongly projecting laterally; fingers very short, with lateral ridges, pointed disks, and very small basal webbings; skin of upper surfaces smooth with a few short weak glandular lines above shoulders and along sides.The best diagnostic characters to recognize Diasporus gularis are its very short hands and feet with pointed fingers and toes, of which the entire terminal disk is dark gray, and the enormous U-shaped gular pouch of the males (Cochran and Goin 1970).Diasporus gularis is most readily distinguished from other Diasporus (i.e., D. anthrax and D. tinker) by its coloration which is orange above with a transverse dark bar between eyes and immaculate venter (dorsum brown with darker blotches and venter dark brown with whitish flecks in D. anthrax; brown above with a tan interorbital bar, dorsolateral flank stripes and venter white with brown marbling in D. tinker ;Lynch 2001).Peters, 1862: Figure 4G Hemiphractus fasciatus Peters 1862: 169.(Linnaeus, 1758) (Duméril & Bibron, 1841) Material examined.Table 1.This species was recently separated from Boana crepitans (Orrico et al. 2017).A large-sized tree frog having its dorsal surfaces pinkish tan with some darker brown spots, blotches or other markings; dark bars on the upper surfaces of the thighs extending onto the rear surfaces of the thighs; ventral surface orange, except for the throat and chest, which are white with some brown flecking.Boana xerophylla may be confused with B. boans and B. pugnax (species distributed in the Magdalena river valley) due to similarities in body size and coloration.However, it may be distinguished by its horizontal pupil and its iris that is pale gray at the center and greenish yellow at the edges (in contrast to a rhomboid pupil with an iris that is black at the center and light orange at the edges in B. boans, and a horizontal pupil with an iris that is black at the center and greenish in the upper portion in B. pugnax).(Cope, 1874): Figure 5B Hyla ebraccata Cope 1874: 69.Dendropsophus ebraccatus -Faivovich et al. 2005: 91.Material examined.Table 1.(Cope, 1886)

Material examined. Table 1.
A small hylid frog with a snout that is short, rounded in dorsal view and slightly rounded in lateral view; head wider than long; skin on dorsal surfaces smooth, with some minute scattered tubercles; axillary membrane present and developed; dorsum reddish brown in females and cream or silver gray in males (Rivera-Correa and Guitérrez-Cárdenas 2012).Dendropsophus norandinus may be distinguished from D. bogerti by having the head wider than long (longer than wide in D. bogerti) and by having finger webbing that extends to the base of the disks of toes I, II, and III (finger webbing absent in D. bogerti), from D. subocularis by lacking light bands connecting eye to edge of lips, and from D. microcephalus by its webbed fingers (fingers free in D. microcephalus).(Dunn, 1934)

Material examined. Table 1.
A small hylid frog with short snout; canthus rostralis very marked; fingers webbed at base about toes I-III; toes webbed nearly to disks of III and V; skin of dorsum smooth and skin of vent rugose (Dunn 1934).Dendropsophus subocularis co-occurs with D. microcephalus and D. norandinus in the Magdalena river valley.It may be distinguished from both by the presence of 2 light bands from the eye to the edge of the lips, enclosing a bar of dark between them (absent in the latter 2 species), and from D. microcephalus by its webbed fingers (fingers free in D. microcephalus).
Our Scinax sp.corresponds to Scinax "A", an undescribed species proposed by Nieto-Castro (1999) using morphological traits, morphometric and chromatic analyses.From that study, the name began to be used to refer to populations of Scinax distributed in Antioquia and Cundinamarca, characterized by having a golden color with a dorsal spotted pattern.(Peters, 1863): Figure 6A Hyla rostrata Peters 1863a: 466.Scinax rostratus -Köhler and Böhme 1996: 139.

Material examined. Table 1.
A moderate-sized hylid frog characterized by a conspicuous acuminate and projecting snout in dorsal view, protruding in lateral view (Duellman 1970).Scinax rostratus is readily distinguished from Scinax ruber and Smilisca sila (both species occurring in the Magdalena river valley) by the combination of its characteristic snout and coloration pattern (surfaces of thighs yellow or pale orange with black or dark brown markings in S. rostratus; distinct black and white markings on posterior thighs in S. ruber; blue spots on flanks and posterior surfaces of the thighs in S. sila).(Laurenti, 1768): Figure 6B Hyla rubra Laurenti 1768: 35.Scinax ruber -Kölher and Böhme 1996: 139.

Material examined. Table 1.
A medium-sized frog with toes webbed except for the innermost; dorsal surfaces generally olive brown or orange-brown; distinct black and white markings on posterior surfaces of thighs.Related species which may be confused with Scinax ruber are S. boulengeri and S. rostratus (also distributed in the Magdalena river valley) from both of these it differs by its smaller size and the presence of black markings on the thighs in form of vermiculations instead of broad and vertical dark bands (Cochran and Goin 1970).Scinax ruber is also similar to Smilisca phaeota from which it differs by lacking green coloration in the rostral region, and to Smilisca sila from which it differs by the absence of blue spots on the flanks and posterior surfaces of the thighs.(Cope, 1862): Figure 6C Hyla phaeota Cope 1862: 358.Smilisca phaeota -Starrett 1960: 303.Material examined.Table 1.(Cope, 1864)    Material examined.Table 1.Heyer, 1994: Figure 6E Leptodactylus colombiensis Heyer 1994: 82.

Material examined. Table 1.
A bicolored caecilian with well-developed tail; lateral stripe yellow, cream or white from head or second collar onto tail; stripe ending at posterior level of vent; subcaudal region dark lilac to blackish (Taylor 1968).Epicrionops parkeri differs from E. bicolor (a species with similar coloration pattern distributed in Colombia) by having a larger body length and 198 primary annuli (in contrast to 117-135 primary annuli in E. bicolor).

Material examined. Table 2.
Adults of this large semiaquatic lizard are easily recognized by their gray to green color and the presence of a single, rounded head crest that is larger in males.Similar species that are found sympatric with B. galeritus are Corytophanes cristatus and Basiliscus basiliscus.The first is usually brownish and has 2 head crests that converge posteriorly.Basiliscus basiliscus has a brown or gray coloration and a single pointed head crest that is larger in males.Males of B. basiliscus also have a large crest in the back that resembles a fin (Maturana 1962;Savage 2002;Köhler 2008).
Anadia gr.ocellata Gray, 1845: Figure 8G Material examined.Table 2.This small lizard may be distinguished from most other mid-to low-elevation gymnophthalmid lizards in having an acuminate snouth and elongate body in combination with smooth head and dorsal scales.Its color pattern includes a series of lateral white spots between the fore and hind limbs (Oftedal 1974, Savage 2002).Usually, anterior most spots are bordered by black, forming ocelli.Although there is no evidence of Anadia ocellata for the Magdalena river valley, we assigned the specimen MHUAR 12694 to Anadia gr.ocellata because its dorsal coloration and 55 transversal rows in contrast to the 44-48 dorsal rows found in A. rhombifera, another species present in the Magdalena river valley (Oftedal 1974, Savage 2002).

Material examined. Table 2.
This species is easily differentiated from other gymnophthalmid lizards in the region by its strongly keeled head and dorsal scales and by having the ventral scales large, keeled, mucronate, and quadrangular to rectangular.It differs from Loxopholis rugiceps, the only other Loxopholis in trans-Andean Colombia, by having the frontonasal divided, instead of single (Köhler 2008).
Material examined.Table 2. Berthold, 1846: Figure 9E Polychrus gutturosus Berthold 1846: 11.This large blunt-headed lizard is immediately recognizable by its extremely long round tail that is at least twice the snouth-vent length, its robust body and relatively thin legs, and the presence of a large subtympanic scale (Peters andDonoso-Barros 1970, Savage 2002).In addition, it lacks toe pads, nuchal, dorsal and caudal crests (Peters andDonoso-Barros 1970, Savage 2002).It differs from Polychrus marmoratus, the other species that could be sympatric, because the latter has a row of    enlarged scales, forming a crest, in the mental and gular regions, which is absent in P. gutturosus (Peters and Donoso-Barros 1970).
Adults have either red or yellow ground color, with a variable dorsal pattern that may consist of black transversal bands or several black dots, mainly located in the posterior region of the scales.Other than coloration, the species differs from other sympatric snakes by having 21-25 dorsal rows at midbody, with the uppermost (3-19) rows keeled (Savage 2002).Some authors recognize an additional species in trans-Andean Colombia, Phrynonax shropshirei (Taylor 1954, Peters et al. 1970, Pérez-Santos and Moreno 1988), but according to Savage (2002), it seems to be a confusion given the wide color variation in P. poecilonotus.Likewise, the profound genetic distinction between P. poecilonotus and P. polylepis (Peters, 1876) revealed by Jadin et al. (2014), who accordingly resurrected the latter species, does apparently not correspond to any morphological distinction.Since neither Jadin et al. (2014) nor the original description (Peters 1876) offer any morphological characters to discriminate between these 2 species, we must assume such distinctions to be obscured by the exuberant variability in Phrynonax for now.Unable to answer the interesting question to which species our specimen actually pertains, at this point we choose a conservative approach and tentatively assign it to P. poecilonotus.(Günther, 1893):  Material examined.Table 2.This small, brown snake has a white, black-bordered line that goes from the postoculars through the fifth and sixth dorsal rows to the tip of the tail.The vertebral region is a little paler than below the lines.Specimens of Rhadinaea decorata often have a subpreocular scale and 110-134 ventral scales (Myers 1974).The most similar species in the region is Urotheca fulviceps, which has a reddish or purple head band, a pale line running along the first and second dorsal scale rows and more than 136 ventrals.(Linnaeus, 1758): Figure 13F Coluber sibon Linnaeus 1758: 222.Sibon nebulatus -Fitzinger 1826: 60. 2. This species has a pattern of gray, white and black bands.Gray bands are larger and with dispersed small black blotches, followed by a thin white band and then by a medium-sized black band (Savage 2002).Ventrally, ground is white with black blotches or bands.Sibon nebulatus has a large head and eyes like species of the genus Imantodes, but it can be easily recognized by its color pattern (Savage 2002).(Daudin, 1803): Figure 13G Coluber compressus Daudin 1803: 247.Siphlophis compressus -Zaher and Prudente 1999: 699.
Trilepida macrolepis (Peters, 1857) Material examined.Table 2.This is a very small species that may be differentiated from other snakes by having body scales of the same size (without enlarged ventral), arranged in 14 rows around de body (Köhler 2008).It has 4 supralabial scales and 10 scale rows around the middle of the tail.Individuals are dark brown dorsally and lighter ventrally, with whitish scale borders (Pinto et al. 2010).Epictia goudotii, a species potentially sympatric with Trilepida macrolepis, has 2 supralabials, a darker color pattern with stripes, and a yellow rostral scale (Pinto et al. 2010).(Garman, 1883): Figure 14E Trigonocephalus asper Garman 1883: 124.Bothrops asper -Hoge 1966: 113.

Discussion
The 43 amphibian species recorded in this study represent 5.2% of the Colombian amphibian species (Frost 2016), and 15% of the species reported from the Cordillera Central in Colombia (IUCN 2016).The 58 reptile species represent 6% of the total Colombian reptiles species (Uetz and Hošek 2016), and 44% of the reptile fauna reported for the Cordillera Central.The Cordillera Central in Colombia is recognized for its high diversity in amphibians and reptiles (Lynch et al. 1997) and the 2 sites we studied represent a high proportion of this herpetofauna.
Fifteen (35%) of the amphibian species found during the monitoring are endemic to Colombia.Most of them are restricted to the eastern flank of the Cordillera Central: Bolitoglossa ramosi, Dendropsophus norandinus, Diasporus anthrax, Epicrionops parkeri, Pristimantis fallax, Pristimantis jaguensis, Pristimantis viejas, Rheobates pseudopalmatus, Rulyrana susatamai, and Sachatamia punctulata.The distribution of the other endemic species extends to the Magdalena river valley (Microcaecilia pricei, Caecilia thompsoni and Craugastor metriosistus), the Chocoan Pacific lowland (Colostethus inguinalis; Grant and Lynch 2004) or the Caribbean lowland and foothills of the Sierra Nevada de Santa Marta (Dendrobates truncatus).Among the reptiles, Anolis sulcifrons was the only recorded species with a distribution restricted to Colombia (Uetz and Hošek 2016).
Five records are of particular interest in this study.Pristimantis jaguensis is an endemic frog from Colombia, with only 1 known locality, the surroundings of the Jaguas hydroelectric project (Rivera-Prieto et al. 2014).Hyalinobatrachium aureogutattum is a frog previously recorded from the departments of Valle del Cauca, Risaralda, Chocó and Antioquia on the western flank of the Cordillera Occidental in Colombia, between 45 and 1570 m above sea level from the provinces of Esmeralda and Imbabura in Ecuador, and from the eastern cordilleras in southwestern Darién province in Panama (IUCN 2016).This is the first record of this species in the Magdalena river valley in Colombia.The presence of Rhadinaea decorata in Colombia was suggested by Myers (1974), and we present a voucher of this species from the San Carlos site, which thus represents the first verified record from Colombia.Anadia ocellata is distributed on both slopes of the cordilleras in Costa Rica and western Panama (Savage 2002).Batista et al. (2016) mention the occurrence in the Pacific lowlands of Colombia, contrary to Oftedal (1974), Savage (2002), and Köhler (2008), who indicated a southern range limit in Panama.We conservatively assign our voucher specimen from the Magdalena river valley to the A. ocellata group (sensu Oftedal 1974).The distribution of Rhinella alata in eastern cordillera was suggested by Pramuck (2006), although her observation lacked a voucher.Here, we present vouchers of R. alata that confirm the inter-Andean distribution of this species with records from Magdalena river valley.
Historically, the herpetofauna from the eastern side of the Andes has been considered very distinct from the trans-Andean region (Duellman 1979).Nevertheless, as part of the long-term sampling carried out in this study, we have recently reported on the occurrence of species distributed in the eastern flank of the northern Cordillera Central but previously known from the Amazon basin: Atractus occipitoalbus (Marin et al. 2017), Anolis fuscoauratus (Grisales-Martínez et al. in press) and Cochranella resplendens (Molina-Zuluaga et al. 2017).These findings urge the need for thorough sampling and a detailed taxonomic revision of species distributed in the Magdalena river valley to elucidate the biogeographic connection between both areas.
We were not able to assign 7 records to any valid species.The record of Marisora sp.(Figure 9F) may correspond to Marisora (Mabuya) candidate species IV suggested by Pinto-Sánchez et al. (2015).The second record refers to an Ameiva sp.In Colombia, there are 3 Ameiva species: Ameiva ameiva, A. bifrontata, and A. praesignis (Harvey et al. 2012).As we only have a specimen photograph (Figure 10B) we cannot assign it to any of these species.The vouchers of Ptychoglossus sp., Sachatamia sp., Colostethus aff.fraterdanieli, and Cercosaura sp. may represent undescribed taxa.
Pristimantis fallax is the only species reported in this study included as Vulnerable at the country level in the Resolucion 192 del 2014 del Ministerio de Ambiente y Desarrollo Sostenible.At the global level, Sachatamia punctulata is listed by the IUCN as Endangered because its extent of occurrence is less than 5000 km 2 , its distribution is severely fragmented, and there is a continuing decline in the extent and quality of its habitat (IUCN 2016).Pristimantis fallax, P. penelopus and Rulyrana susatamai are listed as Vulnerable because of continuous decline in the extent and quality of their habitat (Castro et al. 2004, IUCN 2016).Lastly, 2 species are categorized as Near Threatened: Hemiphractus fasciatus Peters, 1862 and Hyalinobatrachium aureoguttatum (Coloma et al. 2008, Solís et al. 2010; Table 3).
Recently, the conservation status of Colombia's reptiles has been assessed and 43 species are now under one of the 3 threat categories (Morales-Betancourt et al. 2015).However, none of them correspond to species present in our study areas.It is well known that reptiles around the world are declining due to a number of causes directly related to human activities (Gibbons et al. 2000).In addition to habitat loss and degradation, another threat is present in Colombia: the cultural view that reptiles, in particular snakes, are dangerous and nasty animals that should be exterminated (Castaño-Mora 2002, Lynch 2012).This view has probably driven several snake species to the brink of extinction at least locally (Lynch 2012).
To build complete species lists for Colombian amphibians and reptiles evokes large challenges for the following reasons: (1) the high levels of species diversity of amphibians and reptiles in the area, (2) the extreme cryptic diversity in several genera (e.g., Pristimantis, Anolis, Atractus), and (3) the lack of well-identified reference collections.However, continued specimen collection throughout different seasons and years in a specific local-ity will give us a better picture of the regional diversity and will impact future studies aiming to understand the origin and maintenance of this rich fauna.
The high species density, high representativeness of the regional biodiversity, and the high levels of amphibian endemism make these 2 private protected areas very important for conservation of the regional herpetofauna.As shown here, compensation areas of large infrastructure projects (i.e., hydroelectric projects) represent a potential opportunity to preserve the already highly degraded ecosystems in the northern Andes of Colombia.

Figures 1 .
Figures 1. Study area showing the location in Colombia and the northern Cordillera Central.On the right, the two localities (Jaguas and San Carlos) are expanded and black dots represent the sampling sites.Colors represent cover type.

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Colostethus aff.fraterdanieli differs from C. fraterdanieli in having a pale ventrolateral stripe (absent in C. fraterdanieli) and a continuous dark band running along the lateral surfaces of body and head (discontinuous in C. fraterdanieli).It may be distinguished from C. inguinalis by having a complete oblique lateral stripe (partial in C. inguinalis) and from C. pratti by lacking pale dorsolateral stripe (present in C. pratti).

Table 1 .
Voucher list of amphibians of Jaguas and San Carlos hydroelectric protected areas with geographic coordinates.

Table 2 .
Voucher list of reptiles of Jaguas and San Carlos hydroelectric compensation areas with geographic coordinates.

Table 4 .
Taxonomic list of reptiles reported in the Jaguas and San Carlos Hydroelectric protected areas.Asterisks indicate species endemic to Colombia.IUCN Conservation status is shown for both: a global assessment and a national assessment (IUCN 2016/Morales-Betancourt et al. 2015).CR= Critically Endangered, EN = Endangered, VU = Vulnerable, NT = Near Threatened, LC = Least Concern, DD = Data Deficient; NE = Not Evaluated.Geographic distribution follows Lynch et al. (1997): A = Chocoan lowland forests, B = Caribbean lowlands and inter-Andean valleys, C = Llanos, D and E = Amazonia, F = Cordillera Occidental, G = Cordillera Central, H = Cordillera Oriental, I = Sierra Nevada de Santa Marta.