Range extension of Pristimantis uisae ( Lynch , 2003 ) ( Anura : Craugastoridae ) in the Cordillera Oriental of Colombia

Pristimantis uisae is a poorly known frog species reported only from three nearby localities in the western slopes of the Cordillera Oriental of Colombia (Boyacá and Santander Departments). In this work, 12 new records for this species are provided, including the first published records of this species for Cundinamarca Department and from the Eastern slopes of the Cordillera Oriental. Additionally, the new localities reported herein increase its known elevation range to 2300−3050 m. Further studies are needed for having a better understanding of the distribution of Pristimantis uisae.


Introduction
Pristimantis uisae (Lynch, 2003) is a poorly known species of frog endemic to the western slopes of the Cordillera Oriental of Colombia (Boyacá and Santander departments).This species was considered by Lynch (2003) as being closely related to 6 other species of Pristimantis [P.angustilineatus (Lynch, 1998), P. baiotis (Lynch, 1998), P. brevifrons (Lynch, 1981), P. boulengeri (Lynch, 1981), P. dorsopictus (Rivero & Serna, 1987), and P. eremitus (Lynch, 1980)] that inhabit the Cordillera Occidental and the Cordillera Central of the Colombian Andes, as well as the northwestern Andes of Ecuador and southwestern Colombia, and which are now considered as part of the Pristimantis boulengeri species group (with the exception of P. eremitus; González-Durán et al. 2017).However, Lynch (2003) did not provide any characteristic that supports the hypothetical close rela-tionship of P. uisae with these other 6 species, and this species was not assigned to the Pristimantis boulengeri species group by González-Durán et al. (2017).Indeed, the phylogenetic relationships of P. uisae has never been evaluated, as it has not been included in any of the recent morphological or molecular-based phylogenies of the genus (e.g.Hedges et al. 2008, Hoyos et al. 2014, Rivera-Correa and Daza 2016).There are no publicly available DNA sequences for this species (no sequences in the Genbank database as of June 2017), and the only known aspects of its morphology are those listed in its original description (Lynch 2003).
At present, P. uisae is known only from 3 publications: its original description (Lynch 2003) and the ecological studies of Gutiérrez-Lamus et al. (2004), andCortés et al. (2008).Gutiérrez-Lamus et al. (2004) found that this was one of the most abundant species of the frog assemblages in their study, being significantly more common in native NOTES ON GEOGRAPHIC DISTRIBUTION forests than in old oak plantations used for reforestation (Quercus humboldtii), while Cortés et al. (2008) found that this species was evenly found in the interior and edge of 2 fragments of oak forests.The studies of both Lynch (2003) and Gutiérrez-Lamus et al. (2004) were based on the type series of P. uisae, all collected in 2002 from 2 nearby localities in the northwestern slopes of the Cordillera Oriental of Colombia in the departments of Boyacá and Santander (Paipa and Charalá municipalities, respectively).On the other hand, the study of Cortés et al. (2008) was based on specimens found in a locality (Reserva Biológica Cachalú, Encino municipality, Santander) very close to the collecting sites of the other specimens.The possibility that this species exhibits a wider range of distribution has thought possible (Lynch 2004), although no new records have been published to date.However, in an unpublished report, Rueda-Almonacid (2010) documented P. uisae for the first time from Department of Cundinamarca but without giving voucher numbers of collected specimens that serve as the basis for his record.Also, the photographs shown by Rueda-Almonacid (2010) of 2 of these specimens do not allow for unambiguous identification.
As a result of a short field trip to Parque Natural Los Tunos, Municipality of San Antonio del Tequendama (Department of Cundinamarca), and the examination of museum specimens of Pristimantis housed on 2 amphibian collections (Instituto de Ciencias Naturales at the Universidad Nacional de Colombia and Museo de Historia Natural ANDES at the Universidad de Los Andes, Bogotá), 12 new localities are added here to the previously known distribution of P. uisae (Fig. 1, Table 1).
Furthermore, examination of the photographs and description of USNM 146691 given by Cochran and Goin (1970: 426-427, pl.53 G-I), collected [near] "Tequen-dama falls, Municipality of San Antonio del Tequendama" (04°34ʹ39.3ʺN, 074°17ʹ48.8ʺW, 2400 m), suggests that it is probably another unreported specimen of P. uisae, as it is tentatively considered herein.This may also be the case for 3 additional specimens (USNM 146692−146694) collected at the same locality of USNM 146691 and which were also reported erroneously as the Central American species P. cruentus (but not illustrated or described) by Cochran and Goin (1970).However, without examining these specimens, a positive identification is not possible at this time, as they could be a different, sympatric species.

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lecting bag (M.Anganoy, pers.comm.), although many eggs remained in the body of the preserved female.Finally, 2 specimens of P. uisae (ICN 34249, ICN 24250) found in the ICN collection were collected in a locality very close to the paratype locality on Santander Department: Municipality of Charalá, km 38 on the Duitama-Charalá road, Hacienda La Sierra (05°57ʹ56ʺ N, 073°11ʹ08ʺ W, 2400 m).

Results
All of the previously mentioned specimens were identified as P. uisae because they exhibit the following morphological characters that are diagnostic of the species (from Lynch 2003): snout pointed in lateral profile with a papilla on its tip, upper eyelid with at least one small conical tubercle, vomerine odontophores (when present, see below) oval and slanted, males with vocal slits and nuptial pads, a series of small tubercles present in the ulna and another one in the outer tarsi (both ulnar and outer tarsal tubercles being either round, subconical or conical, and even indistinct for the case of the outer tarsus of some specimens), a small round to conical heel tubercle (although in a few specimens it is only poorly defined), rounded subarticular tubercles, and a brown dorsum with darker markings, including labial bars beneath the eyes, as well as limb bars (Figs 2-5).Additionally, while examining the reported specimens it was noted that some specimens (including some paratypes) have a tubercle in the interorbital region, as well as one on each shoulder.These tubercles range from being absent to being conical (Figs 2-5), and they do not appear to be related to locality or sex.However, it was found that in general adult males and juveniles tend to be more tuberculate than females (Figs 2-5), and also that the degree of tuberculation is highly dependent upon the status of preservation of the ethanol-preserved specimens.
Furthermore, vomerine odontophores are lacking in many specimens, including most juveniles, irrespectively of their sex or their locality.Finding interspecific variability in the presence of vomerine odontophores in adults of P. uisae is unusual, as there are few Pristimantis species exhibiting this condition (G.González-Durán, pers.comm.).Further studies are needed in order to determine the reasons lying behind this variability, including the possibility that there are more than 1 species hiding under the name P. uisae.
The identification of most specimens was corroborated by Sandy Arroyo and John D. Lynch, experts on Pristimantis taxonomy at the Instituto de Ciencias Naturales, Universidad Nacional de Colombia.Additionally, the author examined and compared (directly or indirectly) the newly reported specimens to the entire type series of P. uisae (housed in the ICN and in the Colección de Anfibios, Museo de Historia Natural, Universidad Industrial de Santander, Bucaramanga, Colombia, UIS-A).

Discussion
The records of Pristimantis uisae reported herein include the first published records of this species from Cundinamarca Department, and they increase its altitudinal range from 2300 to 3050 m.The range of P. uisae is also extended approximately 272 km to the southwest (in a straight line) from this species' type locality, the northernmost known place where this species have been reported.As P. uisae was known only from 3 nearby localities, the records reported herein show that the species is much more widely distributed than previously thought (Fig. 1).All newly reported localities occur in well-preserved (at least at the time of collecting) humid montane forests.
These new records show that the scarcity of information of P. uisae since its original description is not due to its possible rarity and limited range, but to the lack of adequate study of the collections made in the localities where it is known or suspected to occur.In addition, the poorly known status of P. uisae is also caused by the paucity of collecting in several areas located between the known sites where this species has been found (e.g., the area near Muzo Municipality).Further collecting, as well as a detailed examination of the available specimens collected on these poorly known areas, is needed for better understanding the geographic range of Pristimantis uisae.Additionally, future studies need to evaluate the relationship of this species with the members of the Pristimantis boulengeri group of González-Durán (2017) and with the geographically close, and phenetically similar P. petersi (Lynch & Duellman, 1980), P. tubernasus (Rivero, 1984), P. bogotensis (Peters, 1863) and P. lynchi (Duellman & Simmons, 1978).

Figure 1 .
Figure1.Map of the known localities of Pristimantis uisae, including the 12 new records reported herein.Each number corresponds to each locality in Table1.Red dots indicate the new reports, whereas the yellow and blue ones indicate the type locality and the locality of the paratypes, respectively.The black dot represents the locality reported byCortés et al. (2008), and the white dot, the unpublished record of Rueda-Almonacid (2010).

Table 1 .
List of all known localities of Pristimantis uisae, sorted by department.The datum was used for all coordinates is WGS84.