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https://doi.org/10.11646/zootaxa.5093.2.5
http://zoobank.org/urn:lsid:zoobank.org:pub:F655E1A4-06F7-47D6-B653-6E754004D370
218 Accepted by M. Rivera-Correa: 3 Dec. 2021; published: 26 Jan. 2022
Article ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Zootaxa 5093 (2): 218–232
https://www.mapress.com/zt/
Copyright © 2022 Magnolia Press
A New Species of Red-eyed frog of the genus Pristimantis (Anura:
Strabomantidae) from the Western Slope of the Cordillera Occidental,
Risaralda, Colombia
SEBASTIÁN DUARTE-MARÍN1*, MANUELA MONTOYA-MARÍN2 & JACKELINE RIVERA-GÓMEZ3
1 Grupo Evolución, Ecología y Conservación (EECO), Programa de Biología, Facultad de Ciencias Básicas y Tecnologías, Universi-
dad del Quindío, Armenia, Colombia.
�
sdm950811@gmail.com; https://orcid.org/0000-0001-6201-7527
2 Grupo de Investigación en Biología de la Conservación y Biotecnología (GIBCBT), Corporación Universitaria de Santa Rosa de
Cabal UNISARC.
�
manuelamontoya210@gmail.com; https://orcid.org/0000-0002-3477-769X
3 Grupo de Ecología y Diversidad de Anfibios y Reptiles, Facultad de Ciencias Exactas y Naturales, Universidad de Caldas, Calle 65
# 26-10, A.A. 275, Manizales, Colombia.
�
Jackeline.rivera.gomez@gmail.com; https://orcid.org/0000-0001-7620-4876
*Corresponding author
Abstract.
We describe a new species of red-eyed frog of the genus Pristimantis from the tropical forest at elevations of 1000–1350
m on western slope of the Cordillera Occidental, department of Risaralda, Colombia. The new species differs from conge-
ners by having dorsum skin shagreened with low tubercles; vocal sac single, median and subgular in males, iris red wine
in females copper red in males, and by presenting black groin coloration with yellow light blotches. In addition, the new
species has concave frontoparietals, an unusual character described in Brachycephaloidea frog species.
Keywords. Andes, frontoparietals, Terrarana, taxonomy, new species
Resumen.
Describimos una nueva especie de rana de ojos rojos del género Pristimantis encontrada en bosque tropical a elevaciones
de 1000–1350 m en la vertiente occidental de la Cordillera Occidental, departamento de Risaralda, Colombia. La nueva
especie se diferencia de sus congéneres por la piel dorsal áspera con tubérculos bajos, saco vocal simple, medio y subgular
en machos, iris rojo vino en hembras e ingle de los muslos negros con manchas amarillas claras. Adicionalmente, la nueva
especie presenta frontoparietales cóncavos, un carácter inusual descrito en especies de ranas Brachycephaloidea.
Palabras clave. Andes, frontoparietales, Terrarana, taxonomía, nueva especie
Introduction
The northern Andes play an important role in the diversification of the neotropical biota (Kattan et al. 2004; Herzog
and Tiessen 2017). The great regional biodiversity and endemism in the northern Andes have been explained by
historical events such as the lifting of mountains, the connection between North America and South America, and
climate changes during the Pleistocene (Gentry 1982; Gregory-Wodzicki 2000). Regarding amphibians, their high-
est diversity is found in northern Andes (Lynch and Duellman 1997; Duellman 1999). This richness of amphibian
species has been related with the orogeny and historical biogeography of the Andes through the creation of barriers
and connections that have resulted in speciation and dispersal events (Lynch and Duellman 1997; Castroviejo-
Fisher et al. 2014; Mendoza et al. 2015). Furthermore, evolutionary history and ecological aspects of such as direct
NEW RED EYES PRISTIMANTIS FROM COLOMBIA Zootaxa 5093 (2) © 2022 Magnolia Press · 219
embryonic development without a tadpole stage facilitate the use of all terrestrial environments that have sufficient
humidity for the survival of eggs and adults (Heinicke et al. 2007; Hutter et al. 2017). The northern Andes exhibits
a high diversity and endemism (Lynch and Duellman 1997), and coincide with being one of regions with a high
level of deforestation and changes in land use due to the growth of the human population (Rueda-Almonacid 1999;
Armenteras et al. 2011). This anthropic intervention, has negatively affected the populations of these amphibian
species mainly due to the loss of habitat (Alford and Richards 1999; Lips et al. 2005a, 2005b; Stuart et al. 2008).
The Ethnic Special Management Area (ESMA) Alto Amurrupá is an area with extraordinary diversity of am-
phibian fauna in the northernmost part of the Cordillera Occidental of the Andes in Colombia (35 spp.) where
endemic and threatened species, such as Oophaga histrionica and Phyllobates bicolor (IAvH 2017; Urrutia and
Moya-Robledo 2018). Although information on amphibians has not been fully reported and several areas remain
unexplored, available data shows high levels of species richness and endemism of amphibians in the region (Lon-
doño and Roa-Cubillos 2018; Urrutia and Moya-Robledo 2018; Brown 2020). Therefore, the ESMA Alto Amur-
rupá a potential area for the discovery of new additional species from unexplored areas. In our recent exploration
at the ESMA we collected specimens of one Pristimantis species which exhibits a combination of character states
that have been not previously noticed. Here we provide morphological evidence that leads us to propose this Pris-
timantis as a new species, discovered from a cloud forest adjacent to a National Natural Park (NNP) Tatamá, in the
Cordillera Occidental of Colombia.
Materials and Methods
Fieldwork. One field trip was carried out on December 27, 2019, in the Ethnic Special Management Area (ESMA)
Alto Amurrupá, Santa Cecilia, municipality of Pueblo Rico, department of Risaralda, Colombia, 5.31444 N, 76.15361
W, WGS84 (Fig. 1). Through the visual encounter survey (VES; Crump and Scott 1994) between 10:00–14:00 and
17:00–22:00, four researchers conducted searches to record amphibian individuals in secondary forests.
Morphological analysis. Specimens were euthanized with 2% lidocaine, fixed in 10% formalin and preserved
in 70% ethanol. Measurements were taken with manual calipers to the nearest 0.1 mm under a stereoscope, as
described by Guayasamin (2004), Guayasamin et al. (2006), Duellman and Lehr (2009): (SVL) snout-vent length;
(RUL) radio-ulna length; (HAL) hand length; (TIL) tibia length; (FL) foot length; (HW) head width; (HL) head
length; (UEW) upper eyelid width; (ED) eye diameter; (END) eye-to-nostril distance; (IND) internarial distance;
(SED) snout-eye distance; (IOD) interorbital distance; (NSD) narinal-snout distance; (ETD) eye-tympanum dis-
tance; (TD) tympanum diameter. Toe lengths I and II were estimated by pressing both toes against each other (Du-
ellman and Lehr 2009; Cuellar-Valencia et al. 2021). Frontoparietals position was based in Lynch (1971), consist
in involved careful peeling back of derm of the head. Color in life was determined based on field notes and digital
photos. Morphological comparisons were directly made with examined specimens and/or literature descriptions and
illustrations. Sexual maturity was determined by presence of the vocal slits and matured testes in males and convo-
luted oviducts and/or eggs in adult females. Terminology is that of Lynch and Duellman (1997) and Duellman and
Lehr (2009). Series type reside in the Colección de Anfibios y Reptiles de la Universidad del Quindío, Programa de
Biología, Universidad del Quindío, Armenia, Colombia (ARUQ) and the Museo de Zoología Universidad de Santa
Rosa de Cabal, Colombia (CUS).
Results
Pristimantis blasi sp. nov.
(Figs. 2–5).
Holotype. ARUQ1291 (Figs. 2–3), adult female collected ESMA Alto Amurrupá, Santa Cecilia, municipality of
Pueblo Rico, department of Risaralda, Colombia (5.31444 N, 76.15361 W, 1119 m a.s.l, WGS84), on December 27,
2019, by Manuela Montoya-Marín, Jackeline Rivera-Gómez and Sebastián Duarte-Marín .
Paratypes. (N=8), ARUQ1025–1029, CUS-A145–147 (Figs. 4, 5), male adults collected with the holotype.
Referenced specimens. One subadult female ARUQ1030, and two subadult males ARUQ1031–1032, collected
with holotype.
DUARTE-MARÍN ET AL.
220 · Zootaxa 5093 (2) © 2022 Magnolia Press
Definition. Pristimantis blasi sp. nov. is diagnosed by the following combination of characters: (1) skin on
dorsum shagreen (with scattered tubercles), becoming finely shagreened laterally and coarsely on the flanks; dor-
solateral folds absent; lateral folds present; ventral skin areolate; discoidal fold present, well anterior to groin. (2)
Tympanic membrane partially translucent; tympanic annulus evident, ovoid, corresponding to 1/3 (30%–45%) of
eye length; supratympanic fold distinguished, which extends from posterior corner of orbit along upper edge of tem-
poral region and curved toward the insertion of the arm. (3) Snout subacuminate in dorsal view, rounded in lateral
profile and lacking of papilla; canthus rostralis straight in dorsal view, rounded in profile; two and three subconicals
postrictal tubercles. (4) IOD wider than upper eyelid; craneal crest absent; frontoparietals concave at the proximal
edge in contact with the exoccipital in lateral view; upper eyelid bearing small low tubercle at the posterior margin.
(5) Choanae small, ovoid; not concealed by palatal shelf of maxillary arch; dentigerous processes of vomers promi-
nent, positioned posterior to level of choanae, moderately separated, each dentigerous process of vomers bearing
5–7 teeth. (6) Males with vocal slits and single, median and subgular vocal sac; double nuptial pads glandular and
white, present on the dorsomedial surface on the base of the thumb. (7) Finger I shorter than II; discs truncated. (8)
Lateral fringes on fingers; palmar tubercle in heart-shape (not divided); thenar tubercle oval, slightly smaller than
palmar tubercle; supernumerary tubercles low, distributed on all fleshy parts of palm; subarticular tubercles low,
with rounded base and larger than supernumerary tubercles, hyperdistal tubercle present. (9) Antebrachial tubercle
present; two and three ulnar tubercles small and subconical, not coalesced. (10) Subconical tubercle on heel and
small on outer edge of tarsus, small inner tarsal fold. (11) Oval inner metatarsal tubercles, representing three times
the size of outer tubercle; supernumerary plantar tubercles restricted to the proximal segments of the digits subar-
ticular tubercles rounded, prominent. (12) Toes with narrow lateral fringes, toe webbing absent; toe III shorter than
toe V; toe III extending to distal edge of the penultimate subarticular tubercle of toe IV; toe V reaching distal edge
of distal subarticular tubercle of toe IV; discs and circumferential grooves present on all toes. (13) Two cloacal tu-
bercles present, anterior to cloaca. (14) Polymorphic dorsal pattern (Fig. 4), dorsum darker brown with longitudinal
stripes or blotches pale brown, labial bars cream, longitudinal stripe from the posterior edge of the eye, covering
30–50% of the tympanum, which extends to the flanks; flanks darker brown with cream, in anterior surface with
some dark blotches; groin, anterior and posterior surfaces of thighs black with yellow light blotches; throat and belly
surfaces with black markings on cream or pale yellow; concealed surfaces of tibia with contrasting pattern consist-
ing of black mottling on cream or yellow pale background; pale band across heel; iris red wine in females, copper
red in males. (15) SVL adult males 19.2–31.0 mm (mean = 22.4 ± SD = 3.93 mm; n = 8) and female 40.0 mm.
FIGURE 1. Geographic location in Colombia showing the type locality of Pristimantis blasi sp. nov. in the western slope of
the Cordillera Occidental. (A) Red dot shows the type locality. (B) Map of the Ethnic Special Management Area (ESMA) Alto
Amurrupá (10823 Ha), Santa Cecilia, municipality of Pueblo Rico, department of Risaralda, Colombia. Map produced using
Arc Map, Google earth.
NEW RED EYES PRISTIMANTIS FROM COLOMBIA Zootaxa 5093 (2) © 2022 Magnolia Press · 221
FIGURE 2. Holotype (ARUQ1291; SVL = 40.0 mm) of Pristimantis blasi sp. nov. in life.
Description of the holotype. An adult female with head wider than body; head width 38.9% of SVL; HL 43.7%
of SVL; snout subacuminated in dorsal view and rounded in lateral profile; snout lacking papilla at its tip. Canthus
rostralis concave in dorsal view, in transverse cross section is rounded; loreal region straight, lips no flared. Anterior
part of nostrils is directed laterally; internarial distance 60.4% of eye-to-nostril distance; eye-to-nostril distance
equal to of eye diameter. Upper eyelid bearing small subconical tubercle at the posterior margin, upper eyelid width
is 92.2% of IOD and eye diameter 118.4% of IOD. Craneal crest absent. Frontoparietals region concave at the proxi-
mal edge in contact with the exoccipital in lateral view. Upper edge of tympanum covered by supratympanic fold,
which extends from posterior corner of orbit along upper edge of temporal region and curved toward the insertion of
the arm; tympanum size is 38.4% of eye diameter; tympanum superficial, ovoid, and partially translucent; tympanic
annulus evident, eye to tympanum distance is 58.9% of tympanum diameter; two postrictal subconical tubercles.
Choanae small, ovoid, not concealed by palatal shelf of maxillary arch. Dentigerous processes of vomers prominent,
positioned posterior to level of choanae, moderately separated, bearing 5 vomerine teeth. Tongue longer than wide,
posterior edge notched, posterior 2/3 not adherent to the floor of the mouth. Skin on dorsum shagreen, with scattered
tubercles, laterally finely shagreen and coarsely on the flanks; dorsolateral folds absent. The groin and throat skin
are smooth. Venter areolate. Discoidal fold well anterior to groin. Upper surface of limbs is smooth, with absent
transversal folds over tibia. Radio-ulna length is 22.5% of SVL and hand length is 144.8% of radio-ulna length.
Forearm on outer edge bearing three cream small subconical ulnar tubercles. Elbow fold present. Palmar tubercle
in heart-shape (not divided), twice wider than the oval thenar tubercle. Supernumerary tubercles low, distributed on
all fleshy parts of palm (among six and seven). Subarticular tubercles are rounded and low. Fingers bearing lateral
fringes and truncate disks with rounded borders. Pad on thumb slightly wider than digit above the pad. The pads on
fingers II-IV broadly wider than digits, the fingers III and IV the disks nearly as large as tympanum, ventral pads on
fingers are broader than long. First finger is shorter than second. Tibia length is 51.0% of SVL; subconical tubercle
on heel, three cream small tubercles on outer edge of tarsus. Small inner tarsal fold. Foot length is 50.0% of SVL.
Metatarsal tubercles are subconical. Inner metatarsal tubercles oval twice as long as wide, 3 times the size of outer
metatarsal rounded conical tubercle. Supernumerary plantar tubercles are low (four), restricted to the proximal seg-
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222 · Zootaxa 5093 (2) © 2022 Magnolia Press
ments of the digits. Subarticular tubercles rounded, prominent. Toes bearing narrow lateral fringes, toes lack web-
bing. The pads on toes wider than digits, the pads on toes size order from highest to lowest is 4>5=3>2>1; pads on
toes III, IV and V twice wider than digits; ventral pads truncate, wider than long. Outer pads on toes slight narrower
than the pads on outer fingers. Tip of V reaching distal edge of distal subarticular tubercle of toe IV, tip of toe III
extending to distal edge of the penultimate subarticular tubercle of toe IV.
Coloration of the holotype. In life, darker brown with irregular longitudinal stripe pale brown dorsally; flanks
darker brown with cream, in anterior surface with some dark blotches. Labial bars, nostrils and longitudinal stripe
from the posterior edge of the eye cream. Groin, anterior and posterior surfaces of thighs black with yellow light
blotches. Throat and belly surfaces with black markings on cream or pale yellow; concealed surfaces of tibia with
contrasting pattern consisting of black mottling on yellow pale background; pale band across heel. Cloacal triangle
absent. Iris red wine with a black transversal band. In ethanol, the patterns remain, but the dorsum turns gray and
ventral surfaces turn cream.
Variation. Measurements and ratios of males and female are shown in Table 1. Some males (ARUQ1025,
1028; CUS-A146) have two small suprascapular tubercles. Male ARUQ1025 have W-shaped scapular fold. Some
males (ARUQ1029; CUS-A145–146) have four ulnar tubercles. The dorsal pattern varies from longitudinal stripes
or blotches (Fig. 4). The throat may be dark with irregular cream blotches (ARUQ1027–1029, 1031–1032; CUS-
A146–147). Groin, anterior and posterior surfaces of thighs black with pale blotches (ARUQ1029, 1031–1032; CUS-
A146–147). Ventrally, finger I, II and concealed surfaces of tibia pale orange with black markings (ARUQ1025,
1027; CUS-A145–146, Fig. 5), Iris copper red in males.
FIGURE 3. Illustration of the morphological characteristics the head (lateral and dorsal view), palmar and foot surface of ho-
lotype (ARUQ1291) Pristimantis blasi sp. nov. Scale bar = 5.0 mm. Illustration by SDM and MMM.
NEW RED EYES PRISTIMANTIS FROM COLOMBIA Zootaxa 5093 (2) © 2022 Magnolia Press · 223
TABLE 1. Morphological variation of the type series of Pristimantis blasi sp nov.
Measurements ARUQ1291 ARUQ1025 ARUQ1026 ARUQ1027 ARUQ1028 ARUQ1029 CUS-A145 CUS-A146 CUS-A147
(Female) (Male) (Male) (Male) (Male) (Male) (Male) (Male) (Male)
Snot-vent length (mm) 40.0 23.4 31.0 21.4 20.2 19.4 24.3 19.2 20.5
Radioulna length (mm) 9.0 4.9 6.3 4.9 4.3 4.3 4.9 4.3 4.6
Hand length (mm) 13.0 7.8 10.5 6.4 5.6 5.8 7.8 6.4 6.3
Foot length (mm) 20.0 11.4 16.0 10.0 10.2 9.9 12.3 9.8 10.7
Head width (mm) 15.6 9.2 12.3 8.1 8.0 7.3 9.3 7.6 7.4
Head length (mm) 17.5 10.6 13.9 9.6 9.9 9.0 10.9 9.1 9.8
Upper eyelid width (mm) 3.9 2.2 2.7 2.0 2.0 2.4 2.5 2.1 2.0
Eye diameter (mm) 5.0 3.4 3.6 2.8 3.2 2.5 4.0 2.7 2.8
Eye to nostril distance 5.0 3.9 4.2 2.8 2.8 2.6 3.2 2.8 2.9
Snout to eye distance (mm) 7.1 4.6 4.6 4.4 4.1 3.8 4.5 3.7 3.8
Interorbital distance (mm) 4.2 3.0 3.4 2.8 2.5 2.7 2.7 2.7 2.2
Narinal snout length (mm) 1.7 1.1 0.9 1.1 1.4 1.1 1.1 1.1 1.2
Eye to tympanum distance (mm) 1.1 0.6 0.7 0.6 0.6 0.6 0.7 0.7 0.9
Tympanum diameter (mm) 2.3 1.4 1.4 1.3 1.4 1.5 1.2 1.4 1.5
Finger I length (mm) 7.3 4.1 4.6 3.9 3.4 3.6 4.4 3.3 3.1
Finger II length (mm) 8.6 4.9 6.8 4.5 4.2 4.4 4.9 3.7 4.0
Tibia length/Snout-vent length (%) 51.0 51.4 54.8 49.7 54.2 50.6 51.3 54.4 51.7
Head width/Snout-vent length (%) 38.9 39.4 39.7 37.7 39.5 37.7 38.3 39.4 36.2
Tympanum diameter/Eye diameter (%) 46.5 42.0 39.4 45.6 44.8 59.4 29.9 51.1 54.9
Foot length/Snout-vent length (%) 50.0 48.5 51.6 47.0 50.4 51.0 50.3 50.8 52.3
Head width/Head length (%) 88.9 87.2 88.4 83.8 80.4 81.8 86.0 82.9 76.0
Eye to nostril distance/Head width (%) 32.0 42.6 33.8 34.5 35.3 36.1 33.8 37.3 38.7
Radioulna length/Snout-vent length (%) 22.5 20.5 20.3 23.1 21.5 22.2 20.3 22.4 22.4
Interorbital distance/Head width (%) 26.8 32.1 27.8 34.2 31.7 37.3 28.4 35.6 29.8
Finger I length/Finger II length (%) 84.1 83.2 67.4 86.8 81.5 81.3 88.1 89.3 78.3
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224 · Zootaxa 5093 (2) © 2022 Magnolia Press
FIGURE 4. Illustration of dorsal patterns of Pristimantis blasi sp. nov. (A) ARUQ1030, (B) ARUQ1026, (C) ARUQ1291, (D)
ARUQ1028 and (E) ARUQ1027. Illustration by MMM.
FIGURE 5. Ventral patterns of Pristimantis blasi sp. nov. (A) CUS-A145 (B) ARUQ1026, (C) ARUQ1291, (D) ARUQ1027
and (E) ARUQ1028.
NEW RED EYES PRISTIMANTIS FROM COLOMBIA Zootaxa 5093 (2) © 2022 Magnolia Press · 225
Distribution and Natural history. Pristimantis blasi inhabits primary and secondary in tropical humid forests
of the western slope of the Cordillera Occidental, from 1000 to 1350 m a.s.l (Fig. 6). The individuals of this species
are nocturnal, being active from 2100 to 2300 h; however, it is possible to find inactive individuals during the day
by removing the fallen leaves. The individuals are found in ferns, giant ear plant (Xanthosoma sp.) and trunks at a
height from 0.1 to 1.5 m. The advertisement call of P. blasi consists in a pulsed note similar as “Ñec” to the human
ear, like P. erythropleura and P. cruentus, unfortunately we were unable to record the vocalizing males. The indi-
viduals of this species generally have a strong smell in their groin, similar to the macerated herbs. When finding and
handling these frogs, no defensive behaviors were observed other than fleeing quickly or staying immobile in the
vegetation when found. Sympatric species include Diasporus quidditus, Oophaga histrionica, Phyllobates bicolor,
Pristimantis orpacobates and P. labiosus.
FIGURE 6. (A) The cloud forest of type locality of Pristimantis blasi sp. nov. (B) Landscape showing the mountains of the
ESMA Alto Amurrupá, Santa Cecilia, Municipality of Pueblo Rico. (C) Illustration of Blas A. Cárdenas holding P. blasi sp. nov.
Photographs by Mario A. Santana-Tobar and SDM and Illustration by MMM.
Conservation status of the new species. Pristimantis blasi was found within ESMA Alto Amurrupá (type lo-
cality) and NNP Tatamá (González-Durán coms. pers.). In these localities there are secondary and primary forests,
and the presence of agriculture, which includes livestock, mining activities and logging. Although P. blasi was not
as abundant, 5 to 10 individuals could be found in a few hours of sampling. We suggest that this species should be
“Endangered” according to the IUCN category, by criterion “B1(a)” (extent of occurrence less than 5000 km² and
a number of localities <5).
Etymology. We proposed the etymology blasi in honor to Blas Antonio Cárdenas for his valuable work in the
conservation of Santa Cecilia biodiversity, for his personal contributions to the ecotourism and the protection of
cloud forests in the municipality of Pueblo Rico, which directly benefited the species we describe here (Fig. 6C).
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226 · Zootaxa 5093 (2) © 2022 Magnolia Press
TABLE 2. Diagnostic morphological characters in Pristimantis blasi sp. nov., and comparisons with other similar taxa.
Species
Form of dentigerous
process of vomers Skin of dorsum Dorsolaterals folds
Tympanic
membrane
Cranial
crests
Vocal slits and
vocal sac Nuptial pads on thumb
P. blasi sp. nov. oval
shagreen with some low
tubercles absent present absent present white
P. erythropleura oval smooth present in some morphs present absent absent white
P. bicolor oval
smooth with some low
tubercles absent
present
absent absent gray
P. penelopus oval finely shagreen present in some morphs present absent absent white
P. ruedai triangular smooth poorly defined present absent present grey or brown
P. orpacobates triangular shagreen absent absent present absent white
P. mars oval shagreen absent present absent absent white
P. aemulatus oval finely granular absent absent absent present white
P. cruentus unknown smooth absent absent absent absent white
Continued
Species
Digital
pads
Lateral fringes
on Fingers
Lateral fringes
on Toes
Upper eyelid
tubercle
Ulnar
tubercles
Outher
tarsal
tubercles Heel tubercle Groin coloration Reference
P. blasi sp.
nov. truncated present present subconical subconical nonconical subconical
thighs black with yellow light
blotches This study
P.
erythropleura truncated narrow present subconical conical absent subconical
yellow with black reticulum in
males, carmine red with reduced
black markings in females Lynch (1992)
P. bicolor truncated present present conical subconical subconical triangular yellow in females, red in males
Rueda and Lynch
(1983)
P. penelopus rounded present present subconical nonconical nonconical conical
black whit large yellow or
orange spot
Lynch and Rueda-
Almonacid (1999)
P. ruedai rounded present present absent subconical absent absent brown with cream spots
Ruíz-Carranza et al.
(1997)
P. orpacobates truncated absent absent subconical nonconical subconical nonconical
dark brown with minute white
spots Lynch et al. (1994)
P. mars truncated broad present absent subconical subconical subconical
pinkish with prominent black
bands
Lynch and Ruíz-
Carranza (1996)
P. aemulatus rounded absent absent absent
only
anthebrachial absent subconical black with white spots
Ruíz-Carranza et al.
(1997)
P. cruentus truncated unknown unknown subconical subconical subconical conical
brown with yellow or orange
blotches Savage (2002)
NEW RED EYES PRISTIMANTIS FROM COLOMBIA Zootaxa 5093 (2) © 2022 Magnolia Press · 227
Comparative diagnosis. Pristimantis blasi sp nov. (Fig. 7A) is most similar to five species inhabiting northern
of Cordillera Occidental (P. aemulatus, P. cruentus, P. mars, P. orpacobates and P. ruedai), and three species of
Cordillera Central and Occidental (P. bicolor, P. erythropleura, and P. penelopus) by having nuptial pad in males,
digital discs expanded, digital pads with circumferential grooves, heel tubercle present, dorsolateral fold present
(present in some morph of P. erythropleura), venter areolate, inner tarsal fold present, cranial crests absent, toe V
reaches distal subaticular tubercle of toe IV and bright flash color in the inguinal region (Fig. 7). It differs from
those species by dorsum skin shagreen with low tubercles (smooth in P. erythropleura, P. bicolor, P. ruedai and P.
cruentus), cranial crest absent (present in P. orpacobates), frontoparietals concave at the proximal edge in contact
with the exoccipital in lateral view, a small subconical on the upper eyelid, vocal slits and vocal sac (absent in P.
cruentus, P. orpacobates and P. mars, P. penelopus), iris red wine in females, and groin of thighs black with yellow
light blotches (Fig. 8). Additional differences among species are summarized in Table 2.
FIGURE 7. Adult male of Pristimantis blasi sp. nov., and comparisons with other similar taxa: (A) P. blasi sp. nov, paratype
(ARUQ1027); (B) P. bicolor JDL14350, SVL 27.4 mm, adult male; (C) P. cruentus, Cerro Tacarcuna, Provincia de Darien,
Panamá, individual not collected; (D) P. erythropleura, Natural Reserve Rincón Santo, municipality of Pijao, department of
Quindío, Colombia, adult female not collected; (E); P. orpacobates, municipality of Pueblo Rico, department of Risaralda,
Colombia, adult male not collected; (F) P. penelopus MHUA-A 12274, SVL 32.0 mm, adult male. Photos: (A and D) SDM; (B)
John D. Lynch; (C and E) Marco Rada and (F) Mauricio Rivera-Correa.
Discussion
In Colombia, the northern part of the Cordillera Occidental is a region of high endemism for the direct-develop-
ing frogs of the genus Pristimantis (Duellman 1988; Ruiz-Carranza et al. 1997). Between 1981 and 2004 several
expeditions were carried out and 25 Pristimantis species from this region were described (Lynch 1981; Lynch and
Ardila-Robayo 2004). Since then, only two additional species from this region has been described recently (Rivera-
Correa and Daza 2016; Amézquita et al. 2019). The reduction of descriptions in this region could be mainly due to
the lack of researchers interested in the taxonomy of these frogs, insufficient recent sampling effort, poor evaluation
of scientific collections, or even all of the above, operating together. For example, at the ESMA Alto Amurrupá,
research has focused on systematic, ecology and conservation of poison frog (family Dendrobatidae; Lötters et al.
1999; Negret et al. 2017; Posso-Terranova and Andrés 2018), with less attention to other taxonomic groups, such as,
for example, the genus Pristimantis. This suggests that future field efforts to obtain specimens and new data could
lead to unexpected taxonomic findings.
DUARTE-MARÍN ET AL.
228 · Zootaxa 5093 (2) © 2022 Magnolia Press
FIGURE 8. Variation in lateral view of groin color of Pristimantis blasi sp. nov., and other similar species: (A) P. blasi sp. nov.,
holotype (ARUQ1291); (B) P. cruentus, Cerro Tacarcuna, Provincia de Darien, Panamá, not collected; (C) P. erythropleura,
Natural Reserve Rincón Santo, municipality of Pijao, department of Quindío, Colombia, adult female, not collected; (D) P.
erythropleura, Natural Reserve La Patasola, municipality of Salento, department of Quindío, adult male, not collected; (E) P.
orpacobates MHUA-A09036, adult male; (F) P. penelopus MHUA-A 12274, SVL 32.0 mm, adult male. Photos: (A, C and D)
SDM; (B and E) Marco Rada and (F) Mauricio Rivera-Correa.
Although the phylogenetic relationships of Pristimantis blasi sp. nov. are still uncertain, the similarity in exter-
nal morphology (e.g. digital pads with circumferential grooves, heel tubercle present, inner tarsal fold present and
bright flash color in the inguinal region) of this species with some members of the P. ridens species group (e.g., P.
bicolor, P. erythropleura, P. cruentus, P. penelopus and P. orpacobates), could a suggest a close relationship with
these species. However, this prediction requires future phylogenetic testing.
Historically, an important osteological feature in the taxonomy of brachycephaloid frogs is cranial crests (Trueb
et al. 1993). Cranial crests are defined as ornamentation resulting from bone proliferation (exostosis) in frontopari-
etals (Lynch 1971; Trueb 1973). In Pristimantis genus, cranial crest may be slight ridges above the orbits (i.e. cranial
crest low; P. glandulosus Boulenger 1880; P. quicato Ospina-Sarria et al. 2011) or heavy crest development (e.g.
P. tribulosus Lynch and Rueda-Almonacid 1997; P. rufoviridis Valencia et al. 2011). In addition, the concave posi-
tion of the frontoparietals may resemble slight ridges (e.g. P. blasi sp. nov., P. ruedai Ruiz-Carranza et al. 1997; P.
tamsitti Cochran and Goin 1970; P. cisnerosi and P. chocoensis Reyes-Puig et al. 2020), but there is no bone growth
or exostosis (Fig. 9). Therefore, an external review of the frontoparietals can generate ambiguous perceptions in the
presence of cranial crests (e.g. P. gryllus Barrio-Amorós et al. 2010). A detailed inspection of the frontoparietals
(e.g., clearing and staining the skeletons) is necessary to avoid ambiguity when referring to the cranial crests and
thereby accurately establish their character states in the genus Pristimantis.
Acknowledgments
We thank Blas Antonio Cárdenas, German Darío Gallego and Ángela Arredondo for their assistance in the field. We
thank Mauricio Rivera-Correa and Jhon Jairo Ospina-Sarria for your invaluable comments in the manuscript. Also,
we Marco Rada, Mauricio Rivera-Correa, John D. Lynch and Mario A. Santana-Tobar for providing photographs
of the comparison species. The collection permit for this study was endorsed by resolution 2004 of September 5,
2016 to the University Corporation of Santa Rosa de Cabal (UNISARC) as issued by the National Environmental
Licensing Authority - ANLA.
NEW RED EYES PRISTIMANTIS FROM COLOMBIA Zootaxa 5093 (2) © 2022 Magnolia Press · 229
FIGURE 9. Cranium of adult female of Pristimantis blasi (ARUQ1291) in dorsal view. Scale bar = 5.0 mm.
References
Alford, R.A. & Richards, S.J. (1999) Global amphibian declines: a problem in applied ecology. Annual Review of Ecology,
Evolution, and Systematics, 30, 133–165.
https://doi.org/10.1146/annurev.ecolsys.30.1.133
Alexander Von Humboldt Institute for Biological Resources Research (2017) Amphibian Collection of the Alexander Von Hum-
boldt Institute. Version 37.0. Available from: https://doi.org/10.15472/dma300 (accessed 24 February 2021)
Amézquita, A., Suárez, G., Palacio-Rodríguez, P., Beltrán, I., Rodríguez López, C., Barrientos, L.S., Daza-R, J.M. & Maz-
ariegos, L. (2019) A new species of Pristimantis (Anura: Craugastoridae) from the cloud forests of Colombian western
Andes. Zootaxa, 4648 (3), 537–548.
https://doi.org/10.11646/zootaxa.4648.3.8
Barrio-Amorós, C.L., Guayasamin, J.M. & Hedges, S.B. (2012) A new minute Andean Pristimantis (Anura: Strabomantidae)
from Venezuela. Phyllomedusa, 11 (2), 83–93.
https://doi.org/10.11606/issn.2316-9079.v11i2p83-93
Bernal, M.H. & Lynch, J.D. (2008) Review and analysis of altitudinal distribution of the Andean anurans in Colombia. Zootaxa,
1826 (1), 1–25.
https://doi.org/10.11646/zootaxa.1826.1.1
Boulenger, G.A. (1880) Reptiles et batraciens recueillis par M. Émile de Ville dan les Andes de l’Équateur. Bulletin de la Société
Zoologique de France, 5, 41–48.
Brown, R. (2020) KUBI Herpetology Collection. Version 31.42. University of Kansas Biodiversity Institute.
DUARTE-MARÍN ET AL.
230 · Zootaxa 5093 (2) © 2022 Magnolia Press
https://doi.org/10.15468/ubdwdc
Castroviejo-Fisher, S., Guayasamin, J.M., Gonzalez-Voyer, A. & Vilà, C. (2014) Neotropical diversification seen through glass-
frogs. Journal of Biogeography, 41 (1), 66–80.
https://doi.org/10.1111/jbi.12208
Cochran, D.M., & Goin, C.J. (1970) Frogs of Colombia. Bulletin of the United States National Museum, 288, 1–655.
https://doi.org/10.5962/bhl.part.6346
Cuellar-Valencia, O.M., Arriaga-Jaramillo, F.G., García-Gómez, I., Ceballos-Castro, I., Bolívar-García, W., Velásquez-Trujillo,
D.A., Ortiz-Baez, A.S. & Ospina-Sarria, J.J. (2021) Two New Species of Pristimantis (Anura: Strabomantidae) from the
Serranía de los Paraguas: A Priority Site for Conservation of Amphibians in Colombia. Herpetologica, 77 (1), 72–84.
https://doi.org/10.1655/Herpetologica-D-20-00003.1
Duellman, W.E. (1988) Patterns of species diversity in anuran amphibians in the american tropics. Annals of the Missouri Bo-
tanical Garden, 75, 79–104.
https://doi.org/10.2307/2399467
Duellman, W.E. (1999) Distribution patterns of amphibians in South America. In: Duellman, W.E. (Ed.), Patterns of distribution
of amphibians. A global perspective. The Johns Hopkins University Press, Baltimore, Maryland, pp. 255–327.
Duellman, W.E. & Lehr, E. (2009) Terrestrial breeding frogs (Strabomantidae) in Peru. NTV Science, Münster, 322 pp.
Frost, D.R. (2021) Amphibian Species of the World: An Online Reference. Version 6.1 American Museum of Natural History,
New York. Available from: http://research.amnh.org/herpetology/amphibia/index.php (accessed 24 February 2021)
Garzione, C.N., Auerbach, D.J., Smith, J.J.S., Rosario, J.J., Passey, B.H., Jordan, T.E. & Eiler, J.M. (2014) Clumped isotope
evidence for diachronous surface cooling of the Altiplano and pulsed surface uplift of the Central Andes. Earth and Plan-
etary Science Letters, 393, 173–181.
https://doi.org/10.1016/j.epsl.2014.02.029
Gentry, A.H. (1982) Neotropical floristic diversity: phytogeographical connections between Central and South America, Pleisto-
cene climatic fluctuations, or an accident of Andean orogeny? Annals of the Missouri Botanical Garden, 69, 557–593.
https://doi.org/10.2307/2399084
Graham, A. (2009) The Andes: a geological overview from a biological perspective. Annals of the Missouri Botanical Garden,
96 (3), 371–385.
https://doi.org/10.3417/2007146
Gregory-Wodzicki, K.M. (2000) Uplift history of the Central and Northern Andes: A review. Geological Society of America
Bulletin, 112 (7), 1091–1105.
https://doi.org/10.1130/0016-7606(2000)112<1091:UHOTCA>2.0.CO;2
Guayasamin, J.M. (2004) A new species of Eleutherodactylus (Anura: Leptodactylidae) from the northwestern lowlands of
Ecuador. Herpetologica, 60 (1), 103–116.
https://doi.org/10.1655/02-106
Guayasamin, J.M., Ron, S.R., Cisneros-Heredia, D.F., Lamar, W. & McCracken, S.F. (2006) A new species of frog of the Eleu-
therodactylus lacrimosus assemblage (Leptodactylidae) from the western Amazon Basin, with comments on the utility of
canopy surveys in lowland rainforest. Herpetologica, 62 (2), 191–202.
https://doi.org/10.1655/05-40.1
Hedges, S.B., Duellman, W.E. & Heinicke, M.P. (2008) New World direct-developing frogs (Anura: Terrarana): molecular phy-
logeny, classification, biogeography, and conservation. Zootaxa, 1737 (1), 1–182.
https://doi.org/10.11646/zootaxa.1737.1.1
Heinicke, M., Duellman, W. & Hedges, B. (2007) Major Caribbean and Central American frog faunas originated by ancient
oceanic dispersal. Proceedings of the National Academy of Sciences, 104 (24), 10092–10097.
https://doi.org/10.1073/pnas.0611051104
Hutter, C.R., Lambert, S.M. & Wiens, J.J. (2017) Rapid Diversification and Time Explain Amphibian Richness at Different
Scales in the Tropical Andes, Earth’s Most Biodiverse Hotspot. The American Naturalist, 190 (6), 829–843.
https://doi.org/10.1086/694319
Kattan, G.H., Franco, P., Rojas, V. & Morales, G. (2004) Biological diversification in a complex region: a spatial analysis of
faunistic diversity and biogeography of the Andes of Colombia. Journal of Biogeography, 31, 1829–1839.
https://doi.org/10.1111/j.1365-2699.2004.01109.x
Herzog, S.K. & Tiessen, H. (2017) Climate change and biodiversity in the tropical Andes. Inter-American Institute for Global
Change Research (IAI) and Scientific Committee on Problems of the Environment (SCOPE), Sao Paulo, 348 pp.
Lips, K.R., Burrowes, P.A., Mendelson, J.R. & Parra-Olea, G. (2005a) Amphibian declines in Latin America: widespread popu-
lation declines, extinctions, and impacts. Biotropica, 37, 163–165.
https://doi.org/10.1111/j.1744-7429.2005.00023.x
Lips, K.R., Burrowes, P.A., Mendelson, J.R. & Parra-Olea, G. (2005b) Amphibian population declines in Latin America: syn-
thesis. Biotropica, 37, 222–226.
https://doi.org/10.1111/j.1744-7429.2005.00029.x
Londoño, E. & Roa Cubillos, M.M. (2018) Biodiversidad del Área de Manejo Especia Alto de Amurrupá. Versión 1.1. CARD-
ER—Corporación Autónoma Regional de Risaralda. Conjunto de datos de la lista de verificación.
https://doi.org/10.15472/ctgz2u
NEW RED EYES PRISTIMANTIS FROM COLOMBIA Zootaxa 5093 (2) © 2022 Magnolia Press · 231
Lötters, S., Glaw, F., Kohler, J. & Castro, F. (1999) On the geographic variation of the advertisement call of Dendrobates his-
trionicus Berthold, 1845 and related forms from north-western South America (Anura: Dendrobatidae). Herpetozoa, 12
(1–2), 23–38.
Lynch, J.D. (1971) Evolutionary relationships, osteology, and zoogeography of leptodactyloid frogs. Miscellaneous publication,
53, 1–238.
Lynch, J.D. (1981) Two new species of Eleutherodactylus from western Colombia (Amphibia: Anura: Leptodactylidae). Oc-
casional Papers of the Museum of Zoology, University of Michigan, 697, 1–12.
Lynch, J.D. (1992) Distribution and variation in a Colombian frog, Eleutherodactylus erythropleura (Amphibia: Leptodactyli-
dae). Studies on neotropical fauna and environment, 27 (4), 211–226.
https://doi.org/10.1080/01650529209360880
Lynch, J.D., Ruíz-Carranza, P.M. & Ardila-Robayo, M.C. (1994) The identities of the Colombian frogs confused with Eleu-
therodactylus latidiscus (Boulenger) (Amphibia: Anura: Leptodactylidae). Natural History Museum, the University of Kan-
sas, 170, 1–42.
Lynch, J.D. & Ruíz-Carranza, P.M. (1996) New sister-species of Eleutherodactylus from the Cordillera Occidental of southwest-
ern Colombia (Amphibia: Salientia: Leptodactylidae). Revista de la Academia Colombiana de Ciencias Exactas, Físicas y
Naturales, 20 (77), 347–363.
Lynch, J.D. & Duellman, W.E. (1997) Frogs of the genus Eleutherodactylus (Leptodactylidae) in Western Ecuador: systematics,
ecology, and biogeography. Lawrence, University of Kansas Printing Service, 23, 1–236.
https://doi.org/10.5962/bhl.title.7951
Lynch, J.D. & Rueda-Almonacid, J.V. (1997) Three new frogs (Eleutherodactylus: Leptodactylidae) from cloud forests in east-
ern Departamento Caldas, Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 21,
131–142.
Lynch, J.D. (1998) New species of Eleutherodactylus from the Cordillera Occidental of western Colombia with a synopsis of
the distributions of species in western Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Na-
turales, 22 (82), 117–148.
Lynch, J.D. (1999) Ranas pequeñas, la geometría de evolución, y la especiación en los Andes colombianos. Revista de la Aca-
demia Colombiana de Ciencias Exactas, Físicas y Naturales, 23 (86), 143–160.
Lynch, J.D. & Rueda-Almonacid, J.V. (1999) New species of frogs from low and moderate elevations from the Caldas transect
of the eastern flank of the Cordillera Central. Revista de la Academia colombiana de Ciencias exactas, físicas y naturales,
23 (87), 307–314.
Lynch, J.D. & Ardila-Robayo, M.C. (2004) A new Colombian frog of the genus Eleutherodactylus from the northern Cordillera
Occidental. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 28, 403–408.
Mendoza, Á.M., Ospina, O.E., Cárdenas-Henao, H. & García-R, J.C. (2015) A likelihood inference of historical biogeography
in the world’s most diverse terrestrial vertebrate genus: Diversification of direct-developing frogs (Craugastoridae: Pristi-
mantis) across the Neotropics. Molecular phylogenetics and evolution, 85, 50–58.
https://doi.org/10.1016/j.ympev.2015.02.001
Negret, P.J., Allan, J., Braczkowski, A., Maron, M. & Watson, J.E. (2017) Need for conservation planning in postconflict Co-
lombia. Conservation Biology, 31 (3), 499–500.
https://doi.org/10.1111/cobi.12935
Ospina-Sarria, J.J., Mendez-Narvaez, J., Burbano-Yandi, C. & Bolívar-G, W. (2011) A new species of Pristimantis (Amphibia:
Craugastoridae) with cranial crests from the Colombian Andes. Zootaxa, 3111, 37–48.
Posso-Terranova, A. & Andrés, J. (2018) Multivariate species boundaries and conservation of harlequin poison frogs. Molecular
Ecology, 27, 3432–3451.
https://doi.org/10.1111/mec.14803
Reyes-Puig, C., Yánez-Muñoz, M.H., Ortega, J.A. & Ron, S.R. (2020) Relaciones filogenéticas del subgénero Hypodictyon
(Anura: Strabomantidae: Pristimantis) con la descripción de tres especies nuevas de la región del Chocó. Revista mexicana
de biodiversidad, 91, 1–38.
https://doi.org/10.22201/ib.20078706e.2020.91.3013
Rivera-Correa, M. & Daza-R, J.M. (2016). Molecular phylogenetics of the Pristimantis lacrimosus species group (Anura: Crau-
gastoridae) with the description of a new species from Colombia. Acta Herpetologica. Firenze, 11, 31–45.
Rivero, J.A. & Serna, M.A. (1987) Tres nuevas especies de Eleutherodactylus (Amphibia, Leptodactylidae) de Antioquia, Co-
lombia. Caribean Journal Science, 23, 368–389.
Rueda-Almonacid, J.V. (1999). Anfibios y reptiles amenazados de extinción en Colombia. Revista de la Academia Colombiana
de Ciencias Exactas, Físicas y Naturales, 23, 475–498.
Ruíz-Carranza, P.M., Lynch, J.D. & Ardila-Robayo, M.C. (1997) Seis nuevas especies de Eleutherodactylus Duméril, Bibron,
1841 (Amphibia: Leptodactylidae) del Norte de la Cordillera Occidental de Colombia. Revista de la Academia Colombiana
de Ciencias Exactas, Físicas y Naturales, 21, 155–174.
Savage, J.M. (2002) The amphibians and reptiles of Costa Rica: a herpetofauna between two continents, between two seas.
University of Chicago press, Chicago, Illinois, 934 pp.
Schubert, C. & Clapperton, C.M. (1990) Quaternary glaciations in the northern Andes (Venezuela, Colombia and Ecuador).
Quaternary Science Reviews, 9 (2–3), 123–135.
DUARTE-MARÍN ET AL.
232 · Zootaxa 5093 (2) © 2022 Magnolia Press
https://doi.org/10.1016/0277-3791(90)90014-2
Stuart, S.N., Hoffmann, M., Chanson, J., Cox, N., Berridge, R., Ramani, P. & Young, B. (2008) Threatened Amphibians of the
World. Lynx Edicions, Barcelona, International Union for the Conservation of Nature, Gland and Conservation Interna-
tional, Arlington, Virginia, 758 pp.
Trueb, L. (1973) Bones, frogs, and evolution. Evolutionary biology of the anurans: contemporary research on major problems.
University of Missouri Press, Columbia, Missouri, 132 pp.
Trueb, L., Hanken, J. & Hall, B.K. (1993) Patterns of cranial diversity among the Lissamphibia. In: Hanken, J. & Hall, B.K.
(Eds.), The Skull. Vol. 2. Patterns of Structural and Systematic Diversity. University of Chicago Press, Chicago, Illinois,
pp. 255–343.
Urrutia, N.S. & Moya-Robledo, J. (2018) Caracterización de anfibios del corredor biológico Cértegui - Yerrecuí - Amurrupá.
Sector Amurrupá. Version 1.4. Instituto de Investigaciones Ambientales del Pacífico John Von Neumann-IIAP, Quibdó.
[program]
Valencia, J.H., Yánez-Muñoz, M.H., Betancourt-Yépez, R., Terán-Valdez, A. & Guayasamin, J.M. (2011) Una llamativa nueva
especie de Pristimantis (Anura: Terrana: Strabomantidae) de las estribaciones noroccidentales de los Andes de Ecuador.
Avances en Ciencias e Ingeníerias, Quito, Sección B, 2 (3), 41–45.
https://doi.org/10.18272/aci.v2i3.43
APPENDIX
APPENDIX 1. Examined specimens
Voucher Species Locality Coordinates Altitude
ICN19376-79 P. aemulatus Quebrada Agudelo, Vereda Calles, PNN Las
Orquideas, municipality of Urrao, department of
Antioquia
6.5127, -76.2733 1480
ICN6190 P. bicolor Virolín, municipality of Charalá, department of
Santander
6.1083, -73.1983 1789
ARUQ073,
83, 93
P. erythropleura Finca Palobizcocho, Vereda La Julia,
municipality of Filandia, department of Quindío
4.6858, -75.6522 1822
ICN13335-42 P. cruentus Vereda Campo Alegre, Campamento Chancos,
municipality of Restrepo, department of Valle del
Cauca
460
ICN30335-36 P. mars Quebrada La Empalada, 13km carretera
Mistrato-San Antonio de Chami, municipality of
Mistrató, department of Risaralda
5.3816, -75.8883 1760
ICN42456,
58-59
P. penelopus Carretera El Codo, 1.7km Samaná-Florencia,
municipality of Samaná, department of Caldas
1500
ICN16472-76 P. orpacobates Km. 16 Nutibara-La Blanquita, municipality of
Frontino, department of Antioquia
6.7986, -76.2508 1960
ICN16464 P. ruedai Km. 23 Nutibara-La Blanquita, municipality of
Frontino, department of Antioquia
6.7816, -76.2655 1430
