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Accepted by J. Padial: 10 Dec. 2015; published: 18 Jan. 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4066 (4): 421
–
437
http://www.mapress.com/j/zt/
Article
421
http://doi.org/10.11646/zootaxa.4066.4.4
http://zoobank.org/urn:lsid:zoobank.org:pub:03DC485E-A9B6-4E5B-93D8-A7639BB4BF22
A new small frog species of the genus Pristimantis (Anura: Craugastoridae) from
the northern paramos of Colombia
GUSTAVO A. GONZÁLEZ-DURÁN
1
Laboratorio de Anfibios, Grupo Cladística Profunda y Biogeografía Histórica, Instituto de Ciencias Naturales, Universidad Nacio-
nal de Colombia. Bogotá. Colombia. E-mail: gustavo.gonzalezdu@gmail.com
Abstract
I describe a new species of a small-sized frog of the genus Pristimantis found in the paramo ecosystem (3700 masl) on
the northern slope of Los Nevados National Park, Cordillera Central, department of Caldas, Colombia. This new species
is assigned to the Pristimantis leptolophus species-group, given that Toe V is much longer than Toe III and extends to the
distal edge of the distal subarticular tubercle on Toe IV. The new species differs from other taxa by its dorsal golden or
yellowish color patterns, the absence of nuptial pads, lateral fringes on its fingers and toes, and the absence of vomerine
odontophores. Discriminant analyses of morphometric characters of females of P. leptolophus, P. uranobates, and the new
species separate the new species by snout-vent length, tibia length, eye diameter, eye-to-nostril distance, tympanum diam-
eter, and length of toe III. Vomer terms frequently used to describe species are reviewed, such as the oblique keels of the
vomer and the different forms of the dentigerous process. Species belonging to the high Andean Pristimantis leptolophus
species-group are allopatric, suggesting vicariant speciation in different areas of the paramos.
Key words: Anura, Craugastoridae, Pristimantis, taxonomy, vomers, oblique keels and odontophores
Introduction
In the last years the taxonomy of the superfamily Brachycephaloidea has undergone important taxonomic changes
(Padial et al. 2014a). Craugastoridae was erected as a family by Hedges et al. (2008), but its phylogenetic position
and the taxonomy of its internal taxa (subfamilies and genera) has gone through subsequent changes (Heinicke et
al. 2009; Padial et al. 2009; Pyron and Wiens, 2011; Ohler and Dubois, 2012; Canedo and Haddad, 2012; Padial et
al. 2014 a and b). The genus Pristimantis was resurrected by Heinicke et al. (2007), and Hedges et al. (2008)
provided a non-functional diagnosis (i.e., the characters cited by the authors do not permit a generic assignation of
species) and no morphological synapomorphies have been proposed so far for the genus (Padial et al. 2014a).
Pristimantis is the most diverse vertebrate genus in the world (ca. 482 spp; Frost, 2015), distributed from Honduras
to Argentina, across the Andes, Amazonian lowlands, Atlantic forest, Guianan highlands, and the Lesser Antilles
(Duellman and Lehr, 2009), with the greatest species diversity occurring in Colombia and Ecuador (Lynch, 2001;
Pinto-Sánchez et al. 2012) with 213 and 181 species, respectively (Frost, 2015).
The Pristimantis leptolophus species group was proposed by Lynch (1991) in order to accommodate five
species from the Central Cordillera of Colombia. The definition of this group is based on their relatively large
digital discs (other species present little or no expansion) and by the fifth toe being longer than the third (the tip of
toe V reaches the distal subarticular tubercle on toe IV; Lynch 1994). These characters, however, have never been
tested as synapomorphies in a phylogenetic context. Posteriorly, Hedges et al. (2008) added the following
diagnostic characters to the P. leptolophus species group: small size, expanded digital pads, an “S” condition of the
abductor muscle, females reaching a maximum size of 30 mm, robust bodies, narrow heads, short snouts, and
moderately long legs, finger I shorter than finger II, cranial crests absent, and vocal slits and vomerine teeth
present.
Currently, the Pristimantis leptolophus species group is composed of seven species which have never been
included in any phylogenetic analysis; six of them occur in the paramo and subparamo ecosystems, along the
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Cordillera Central of Colombia [P. leptolophus (Lynch, 1980), P. maculosus (Lynch, 1991), P. parectatus (Lynch &
Rueda-Almonacid, 1998), P. peraticus (Lynch, 1980), P. scoloblepharus (Lynch, 1991), P. uranobates (Lynch,
1991)], and one in a northern paramo of the Cordillera Occidental—P. lasalleorum (Lynch, 1995). During
fieldwork on the Cordillera Central of Colombia in the department of Caldas, I found specimens of Pristimantis in
the paramo ecosystem that share the characteristics described for the P. leptolophus species group. Herein, I
describe and name this newly discovered population as a new species.
Materials and methods
Specimens were fixed in 10% formalin and preserved in 70% ethanol. Acronyms are: Instituto de Ciencias
Naturales, Universidad Nacional de Colombia, Bogotá, Colombia (ICN), The University of Kansas Natural History
Museum and Biodiversity Research Center, Kansas, EEUU (KU), Field Museum of Natural History, Chicago,
EEUU (FMNH).
Type material of Pristimantis uranobates, P. maculosus, P. scoloblepharus, and P. parectatus, were personally
revised, as well as photographs and drawings of the holotypes and new specimens of P. lasalleorum, P.
leptolophus, and P. peraticus collected near the type localities. Sexual maturity was determined by presence of the
vocal slits and matured testes in males and convoluted oviducts or eggs in adult females. Terminology is that of
Lynch and Duellman (1997) and Duellman and Lehr (2009).
Techniques for cleared and double stained specimens were taken from Taylor and Van Dyke (1985), the
osteological drawings and descriptions were made from adult females (ICN-55591-92, 55697), following the
osteological terminology of Duellman and Trueb (1986) and Trueb (1973).
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): (1) snout-vent length; (2) radio-ulna
length; (3) hand length; (4) tibia length; (5) foot length; (6) head width; (7) head length; (8) upper eyelid width; (9)
eye diameter; (10) eye-to-nostril distance; (11) internarial distance; (12) snout-eye distance; (13) interorbital
distance; (14) narinal-snout length; (15) eye-tympanum distance; (16) tympanum diameter; (17) finger I length
(from base of subarticular tubercle to fingertip); (18) Finger II length (from base of subarticular tubercle to
fingertip).
Discriminant analyses were employed to assess the morphometric distinctiveness between the new species
(n=17) and P. leptolophus (n=21) and P. uranobates (n=22)—the most phenotypically similar ones. Considering
that there are problems in differentiating between males of certain species and that the major morphometric
differences in Pristimantis occur between females, only adult females were used for this analysis. In addition to the
18 morphometric variables (see above), the length of fingers III, IV and toe lengths III, IV, V (from the base of
subarticular tubercle to fingertip), and the width of fingers discs III and IV, and width of toes discs IV and V were
also included in the discriminant analysis.
All statistical analyses were performed on Statistica 7 software. For the discriminant function analysis, scores
were projected in the space of the canonical axis, allowing a graphical differentiation of the species analyzed,
according to the forward stepwise method. Wilks's lambda distribution and coefficients for the roots were used to
determine which characters are most important in the morphometric differentiation. Variables which most
accounted for the discriminant analysis were tested with an analysis of variance (ANOVA) to compare means, and
Tukey’s HDS multiple comparisons test was used when means were considered not homogeneous.
Pristimantis stictus
sp. nov.
(Figures 1–2)
Holotype. ICN 55689, adult female collected at the vereda el Vergel, Municipality of Marulanda, Department of
Caldas, Colombia (5.2372 N, 75.3512 W; 3700 m.a.s.l), by Gustavo Gonzalez-Duran, on 30 January 2010.
Paratypes. Nine adult females ICN-55690-92, 55696-99, 55702, 55704 and eight adult males ICN-55693-95,
55700-01, 55703, 55705-06, collected with the holotype on 29–30 January 2010. Six adult females ICN-55710,
55712-16 and four adult males ICN-55707-09, 55711, collected on 3–4 November 2012, in San Pablo,
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Municipality of Neira, Department of Caldas, Colombia (5.1193 N, 75.3234 W; 3700 m), by Gustavo Gonzalez-
Durán.
FIGURE 1. A. Ventral view of Pristimantis stictus sp. nov., adult female holotype, (ICN-55689). B. Dorsal view of P. stictus
sp. nov., adult female holotype, (ICN-55689). C. Ventral view of hand of P. stictus sp. nov., adult female holotype, (ICN-
55689). D. Ventral view of foot of P. stictus sp. nov., adult female holotype, (ICN-55689). E. Lateral view of P. stictus sp. nov.,
adult female holotype (ICN-55689). White bars equal 4 mm.
Referred specimens. ICN-14417 one adult male and ICN-14418-23, 14432 eight adult females collected on
23 June 1984 in the vicinity of the Hotel Termales del Ruiz, Municipality of Villamaria, Department of Caldas,
Colombia (4.9711 N, 75.3844 W; 3370 m). ICN-55591-92 adult female collected on 28–29 November 2011 vereda
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El Desquite, Municipality of Manizales, Department of Caldas, Colombia (5.0650 N, 75.3716 W; 3750 m). ICN-
55717 adult female collected on 25 November 2007 in the vereda Hojas Anchas, Municipality of Salamina,
Department of Caldas, Colombia (5.2423 N, 75.3702 W; 3670 m). ICN-55718-31 collected on 1–5 March 2014
vereda El Desquite, Municipality of Manizales, Department of Caldas, Colombia (5.0651 N, 75.3791 W; 3620 m).
ICN-55732-33 juveniles collected on 30 January 2010 in the vereda el Vergel, Municipality of Marulanda,
Department of Caldas, Colombia (5.2372 N, 75.3512 W; 3700 m). ICN-55734-39 six juveniles and ICN-55741 one
adult male collected on 3–4 November 2012, in San Pablo, Municipality of Neira, Department of Caldas, Colombia
(5.1193 N, 75.3234 W; 3700 m).
Etymology. The specific epithet stictus is derived from greek stiktos (spotted) due to the spots on the groin and
belly. The name is used as an adjective.
FIGURE 2. A. Lateral aspect of Pristimantis stictus sp. nov., adult male (ICN-55740). B. Lateral view of P. stictus sp. nov.,
adult female (ICN55715). C. Lateral view of P. stictus sp. nov., adult female (ICN-55713). D. Lateral aspect of P. stictus sp.
nov., adult female (ICN-55716). E. Ventral view of P. stictus sp. nov., adult female (ICN-55713). F. Ventral view of P. stictus
sp. nov., adult female (ICN-55716).
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FIGURE 3. Variation in life coloration. A. Pristimantis leptolophus, adult female (ICN-7799) (Photo: John D. Lynch); B.
Pristimantis uranobates, adult female (ICN-55588); C. Pristimantis peraticus, adult female (ICN-55742) (Photo: Marco Rada);
D. Pristimantis maculosus, adult female (ICN-55578); E. Pristimantis parectatus, adult female (ICN-55743) (Photo: Marco
Rada); F. Pristimantis scoloblepharus, holotype adult female (ICN-8583) (Photo: John D. Lynch).
Generic and familial allocation. Despite the absence of phenotypic synapomorphies for Pristimantis (see
Padial et al. 2014a,b), the new species is assigned to this genus based on its similarity to other species in this genus
occurring in the area, as well as by characters listed by Hedges et al. (2008), such as the absence of cranial crests,
expanded terminal discs on digits, well-defined circumferential grooves with T-shaped terminal phalanges, and
non-protruding subarticular tubercles. The new species is assigned to the P. leptolophus group due to females with
a maximum SVL of less than 30 mm, narrow heads, short snout, Finger I shorter than Finger II, and Toe V much
longer than Toe III and it extends to the distal edge of the distal subarticular tubercle on Toe IV. The position of the
new species should be evaluated through future phylogenetic analyses.
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TABLE 1. Diagnostic morphological characters in the Pristimantis leptolophus species group.
continued.
continued.
SVL males SVL females Dorsal coloration in life Groin coloration in
life
Dorsolaterals folds
P. lasalleorum 15.6 18.4–21,0 Dark brown above Brown Low
P. leptolophus 14.3–18.2 21.2–25.1 Brown with darker
markings
Brown Low and thin
P. maculosus 17.8–19.2 23.8–26.3 Dark brown Black with large
white spots
Inconspicuous, reach
anterior to level of
sacrum
P. parectatus 14.2–16.0 22.5–27.0 Brown with darker
markings
Pale brown or
cream
Low and thin
P. peraticus 15.3–18.3 20.7–26.0 Pale Brown Brown Low and thin
P. scoloblepharus 17.4–20,3 23.7 Brown above with dark
brown mottling, flanks
paler
Yellow spots Thickened with conical
warts
P. uranobates 15.7–19.8 23.1–27.8 Brown with markings
(polymorphic pattern)
Brown spots and
reticulation
Thickened,
polymorphic coloration
Lateral fringes
on finger
Nuptial pads
on thumb
Ulnar
tubercles
Tarsal
tubercles
Heel tubercle Tubercles on
upper ey elid
P. lasalleorum Present Large Subconical Small Present,
subconicals
Conical
P. leptolophus Narrow Absent Elongate Conical Conical Present, small
P. maculosus Lateral keels Present Small Small Present, small Small warts
P. parectatus Absent Absent Subconical Subconicals Present,
subconicals
Short conical
P. peraticus Narrow Absent Elongate Small Small subconicals Present, small
P. scoloblepharus Narrow Present Present Small Conicals Present, conicals
P. uranobates Narrow Present Elongate Subconicals Present,
subconicals
Present,
subconicals
Snout Tympanum Cranial
crests
Vomerine
odontophores
Source
P. lasalleorum Short, rounded in dorsal and
lateral view
Present Absent Absent Lynch, 1995, this work.
P. leptolophus Short, rounded in dorsal and
lateral view
Present Absent Absent Lynch, 1980, Lynch,
1991.
P. maculosus Short, subacuminate in dorsal
and rounded in lateral view
Present Absent Small, low Lynch, 1991, this work.
P. parectatus Short, subacuminate in dorsal
and rounded in lateral view
Present Absent Low, oval Lynch & Rueda-
Almonacid, 1998.
P. peraticus Short, subacuminate in dorsal
and rounded in lateral profile
Absent Low Low Lynch, 1980, this work
P. scoloblepharus Subacuminate in dorsal and
rounded in lateral view
Present Absent Obliques Lynch, 1991.
P. uranobates Round in dorsal and lateral view Present Absent Obliques Lynch, 1991, this work
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FIGURE 4. Variation in lateral view of groin color of Pristimantis stictus sp. nov., and comparisons with other similar taxa. A.
P. stictus sp. nov., adult female (ICN-55702); B. P. parectatus, adult female (ICN-55740); C. P. maculosus, adult female (ICN-
55578); D. P. scoloblepharus, adult females (ICN-55744) (Photo: Marco Rada), SVL: 26.5 mm; E. P. uranobates, female not
collected; F. P. peraticus, female not collected (Photo: Marco Rada).
Diagnosis. The new species can be diagnosed by the following combination of characters: (1) Dorsal skin
smooth; dorsolateral folds present, continuous from eyelid to the sacrum; ventral skin coarsely areolate; discoidal
fold absent. (2) Tympanum superficial, tympanic annulus prominent, round, corresponding to 1/2–1/3 (35%–46%)
of eye length, with an obscure stripe on the supratympanic fold. (3) Snout short, subacuminate in dorsal view,
rounded in lateral profile; canthus rostralis concave. (4) IOD wider than upper eyelid; craneal crests absent, upper
eyelid bearing small subconical tubercle. (5) Vomerine odontophores absent. (6) Males with vocal slits and
subgular vocal sac; nuptial pads in males absent. (7) Finger I shorter than II, with large rounded digital disc. (8)
Lateral fringes on fingers. (9) Ulnar tubercles small, subconical. (10) Subconical tubercles on heel and outer edge
of tarsus, small inner tarsal fold. (11) Two oval inner metatarsal tubercles, representing six times the size of outer
tubercle; supernumerary plantar tubercles present. (12) Toes with lateral fringes; toe discs slightly narrower than
those on fingers. (13) Dorsum brown, yellowish or copper, with golden tones, in some cases spotted yellow;
laterally light brown; groin yellowish with dark spots; venter and legs yellow with dark spots. (14) Adults small,
males 17.8–22.2 ( = 19.9 ± 1.2; n=15) and females 22.0–28.2 ( = 25.4 ± 1.6; n=17).
x
x
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Comparisons. Pristimantis stictus differs from other species of the cloud forest from the northern of
Cordillera Central (P. maculosus, P. parectatus, and P. scoloblepharus) by the absence of vomerine odontophores,
low and thin dorsolateral folds extended from the posterior eyelid to beyond the sacrum, small tubercles on the
eyelid, and by lacking nuptial pads (Figure 3). Pristimantis stictus differs from similar paramo species (P.
lasalleorum, P. leptolophus and P. peraticus) by presenting tympanic annulus and cavum tympanicum, more
enlarged discs, and broad lateral fringes on fingers (Figure 3). It can be easily distinguished from the sympatric
species P. uranobates by the absence of vomerine teeth and nuptial pads, and yellow groin with black reticulations
(Figure 4). Additional differences among species are summarized in Table 1.
Description of the holotype (Figure 1). Head wider than body; snout rounded in dorsal view and in lateral
profile. Snout short, canthus rostralis concave, loreal region is slightly concave, sloping abruptly to lips; lips not
flared. Nostrils protruding, directed laterally. Upper eyelid bearing one small subconical tubercle. No cranial crests.
Supratympanic fold thin, obscure on upper and posterior edges of tympanum. Tympanum superficial, round,
separated from eye by a distance equal to its size, tympanic annulus prominent. Postrictal tubercles small,
subconical. Choanae small, round, not concealed by palatal shelf of maxillary arch. Vomerine odontophores absent,
represented by oblique keels posteromedial to choanae, not bearing teeth. Tongue longer than wide, posterior edge
not notched, posterior one-half not adherent to floor of mouth. Skin of dorsum finely shagreen with small and low
spicules. Dorsolateral folds extending from eye to sacral region, lacking thickening or distinctive coloration. Flanks
granular, more distinctive than dorsum, with numerous low warts. Groin smooth. Tympanic folds with
superposition of tubercles, especially posterior to tympanum. Ventral surface areolate and throat weakly areolate.
Discoidal fold indistinct. Upper surface of limbs smooth, without ridges. No cloacal sheath. Two postcloacal warts.
Forearm bearing a row of three subconical ulnar tubercles. A broad lateral fringe along outer edge of palm and
continuing along outer edge on finger IV. Palmar tubercle bifid, wider than oval tenar tubercle. Six supernumerary
tubercles. Subarticular tubercles rounded and nonconical. Fingers bearing broad lateral fringes, and round disks.
Pad on thumb slightly wider than digit proximal to pad. The pads on fingers II–IV wider than digits, those of
fingers III and IV have discs as large as tympanum or twice the width of digit; ventral pads, broader than long. First
finger shorter than second. Subconical tubercle on heel, two small tubercles on outer edge of tarsus. Inner tarsal
fold present, its length equals one third of tarsus. Inner metatarsal tubercule twice as long as wide, six times the size
of the rounded subconical outer tubercle. Plantar surfaces bearing numerous low supernumerary tubercles.
Subarticular tubercles rounded, nonconical. Toes bearing lateral fringes, narrower than fringes on the fingers; toes
lack webbing. Discs on toes III, IV and V wider than digits; discs on toes IV and V wider than those of the inner
toes; ventral pad as wide as long. Outer pads on toes narrower than the pads on outer fingers. Tip of toe V reaches
proximal third of distal subarticular tubercle of toe IV, tip of toe III reaches the distal border of penultimate
subarticular tubercle of toe IV. Heels broadly overlapping when hindlimbs are flexed perpendicular to the sagittal
plane.
Measurements of the holotype (in mm). Snout-vent length =26.7, radio-ulna length=6.3, hand length=8.3,
tibia length=12.5, foot length=12.7, head length=10.1, head width=8.9, upper eyelid width=2.3, eye diameter=3.2,
eye to nostril distance=2.2, internarial distance=2.4, snout to eye distance=4.3, interorbital distance=3.5, eye to
tympanum distance=0.8, tympanum diameter=1.5, finger I length=2.2, finger II length=3.0.
Coloration of the holotype. In life, copper-colored or brown dorsally; laterally and dorsal surfaces of limbs
have a lighter brown color. The groin presentes a yellowish color with elongated black spots. The belly is bright
yellow with small black dots. The throat is copper. The undersides of thighs have small black dots, while the spots
are elongated on the posterodorsal surface. Cloacal triangle absent. Dorsolateral folds lacking a distinctive color.
Iris yellow with a reddish transversal band. Rostral markings inconspicuous. In ethanol, the patterns remain, but the
dorsum turns gray and ventral surfaces turn cream.
Adult cranial osteology (Figure 5). The maximum width of the skull is at the posterior level of the maxillae.
Bones of the skull are well ossified. Both centers of ossification of the sphenethmoid are fused dorso and
ventromedially; dorsally, it slightly overlaps the anterior part of frontoparietals and a small portion of the
posterolateral border of the nasals. The nasals are medially separated; the maxillary process does not reach the
preorbital process of the maxilla. Nasals do not articulate with the frontoparietals. The frontoparietal fenestra is
absent. No cranial crest. Posterolaterally frontoparietals extend to the level of the epiotic eminence of the prootic.
The frontoparietals are not fused with the prootic. The sphenethmoid ventrally subtends the dentigerous processes
of vomers, and the cultriform process of parasphenoid. The cultriform process reaches anterior to the level of
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palatines. Dorsally, the otoccipital bears a cartilaginous parotic crista that connects with the otic ramus of the
squamosal. Stapes present. The vomers distinctly medially separated, prechoanal process indistinct; the
postchoanal process is directed laterally and narrow. Dentigerous processes of vomers are elongated and narrow,
directed posteromedially, extending anteriorly to palatines; externally, it is represented by oblique keel of vomer
(Figure 5C–D). Vomerine teeth are absent.
FIGURE 5. Cranium of adult female of P. stictus (ICN-55591; SVL: 25.6 mm) in dorsal (A) and ventral view (B), bars
represent measurements of 2.5 mm, black indicates foramina, gray cartilage and stippling bones; C. Photography of oblique
keel of vomer slightly concealed by mucosa in P. stictus (ICN-55697; paratype); D. Vomer in cleared and stained specimen of
adult female of P. stictus (ICN-55692).
Va ri at io n . Measurements and ratios of males and females are shown in Table 2. The head is wider than the
body in males and narrower or equal in females. The snout is round in lateral profile in females and in some it may
be slightly protruded (ICN-55701, 55707-08). Males and some females (ICN-55691, 55693-95, 55697, 55701,
55703, 55707-709, 55711, 55714-16) have a tubercle on the tip of the snout. The supratympanic stripe is not
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evident in some females, but it is present in all males and some females (ICN-55690, 55693, 55695, 55697, 55702,
55713). All females lack vomerine odontophores, however, some (ICN-55692-93, 55696-97, 55699, 55701-02,
55704, 55712) possess the oblique keel that lies posterior to the choanae. Some males (ICN-55705, 55708-09,
55711) do not present the oblique keel while others (ICN-55698, 55707) may have a tooth in the posterior end of
the oblique keel. The dorsal skin color varies from dark brown to yellowish or copper. The groin may be yellow in
females, with small dark dots or elongated spots (ICN-55690, 55692, 55696, 55713). Males usually possess spots
on groin (ICN-55694, 55709, 55711, 55714). The belly is yellowish with black spots in females, while males do
not present such pattern (ICN-55694, 55705, 55708-09, 55711). The gular region of males is dark grey. Females
present small black dots on the ventral part of thighs and elongated spots on posterodorsal areas. Dorsolateral folds
have no distinctive color. The iris varies from reddish to yellow.
TABLE 2. Measurements and morphological proportions (in percentages) of adult females and males of Pristimantis
stictus (range, mean, and standard deviation (SD) in mm). Head width (HW); snout-vent length (SVL).
Females n=17 Males n=15
Character Range Mean SD Range Mean SD
Snout-vent length 22–28.2 25.4 1.6 17.8–22.2 19.9 1.2
Radioulna length 5.8–7.4 6.4 0.4 4.6–5.7 5.1 0.3
Hand length 7.4–9.4 8.4 0.5 5.8–7.2 6.5 0.4
Tibia length 11.8–13.8 12.9 0.6 9.3–11.2 10.1 0.5
Foot length 11.7–14.9 13.2 0.9 8.8–11.4 10.2 0.8
Head width 8.6–10.2 9.6 0.5 6.8–8.6 7.6 0.5
Head length 7.7–9.6 8.4 0.5 6.2–7.4 6.9 0.4
Upper eyelid width 1.8–2.5 2.1 0.2 1.6–2.2 1.8 0.2
Eye diameter 2.7–3.7 3.2 0.3 2.2–2.9 2.2 0.2
Eye to nostril distance 2–2.6 2.3 0.1 1.3–2.1 1.2 0.2
Internarial distance 2.2–2.8 2.6 0.2 1.8–2.3 2.1 0.2
Snout to eye distance 3.5–4.3 4.0 0.2 2.9–3.5 3.2 0.2
Interorbital distance 3–3.9 3.3 0.2 2.1–3 2.6 0.2
Narinal snout length 1.3–1.9 1.5 0.1 1.1–1.4 1.3 0.1
Eye to tympanum distance 0.6–1.2 0.8 0.2 0.4–0.8 0.7 0.1
Tympanum diameter 1.1–1.6 1.4 0.1 0.9–1.3 1.1 0.1
Finger I length 1.8–2.6 2.2 0.2 1.3–1.9 1.6 0.2
Finger II length 2.5–3.5 2.9 0.3 1.9–2.6 2.3 0.2
Toe III length 3.9–5.1 4.4 0.3 – – –
Toe IV length 6.5–8 7.3 0.6 – – –
Tibia length/SVL 47.6–55.5 50.8 2.1 46.9–54.5 51.1 2.7
Head width/SVL 35.1–41,9 37.9 1.4 36.1–40.7 38.5 2.1
Tympanum diameter/Eye diameter 31.0–47.7 46.7 5.4 35.9–56.2 41.6 5.2
Foot length/SVL 46.0–57.5 51.1 2.2 47.4–53.4 51.2 3.1
Head width/head length 102.7–123.4 114.3 4.4 105.6–124.2 111.1 4.9
Eye to nostril distance/HW 19.7–25.3 24.0 1.8 21.7–26.9 23.1 2.0
Radioulna length/SVL 23.6–28.7 25.9 1.2 23.5–27.8 25.7 1.4
Interorbital distance /HW 25.7–38.9 34.7 2.1 30.7–37.6 33.8 2.9
Finger I length/Finger II length 60.8–82.6 74.5 5.2 64.3–82.1 70.7 5.4
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FIGURE 6. Map of Colombia showing the distribution of Pristimantis stictus sp. nov. and other species. Black circles indicate
localities of P. stictus sp. nov., and a white circle signals the type locality. Distribution of other species in the P. leptolophus
group: P. maculosus (black five pointed star), P. parectatus (white diamond), P. lasalleorum (inverted triangle), P. uranobates
(white four pointed star), P. peraticus (black six pointed star), P. scoloblepharus (black triangle), and P. leptolophus (white
squares).
Distribution and natural history. The new species is found at altitudes above 3500 m, reaching a maximum
altitude of 3700 meters. At altitudes of 3500 to 3600 m, P. stictus is parapatric with P. uranobates. So far, P. stictus
is endemic to the department of Caldas, in the northern region of Páramo Los Nevados (Figure 6), being very
abundant throughout its distribution range. It inhabits high altitude vegetation of the high Andean forest, and
Subparamo (Rodríguez et al. 2006). Pristimantis stictus was found actively at night mainly associated with
terrestrial bromeliads. Males called at heights lower than 70 cm. Amplectant pairs and gravid females were found
on January 2010 and April 2014. Juveniles were found on January 2009 and November 2012.
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Morphometry between similar species. Discriminant analysis allowed to differentiate species. Variables that
showed significant differences between means are listed in Table 3. The variables indicating a higher proportion of
variance were toe III length, snout-eye distance, snout-vent length, Toe IV length, and eye-to-nostril distance
(Table 3). The Chi-square Test with successive roots removal showed significant association between the species
and canonical functions for the first (R= 0.95; X
2
= 170.09; df= 44 p <0.00) and second root (R= 0.85; X
2
= 57.87;
df= 21 p <0.00). Such variation allowed the recognition of three distinct groups corresponding to the target species
(Table 3 and Figure 7).
TABLE 3. Variables in the model for the Discriminant Function Analysis summary and standardized coefficients for the
two roots. Wilks’ Lambda: 0.02578, F(44,72)=8.5544, P<0.0000. Abbreviations: Eigenvalue (Eig), cumulative
proportion (Cp).
Pristimantis stictus differs from P. leptolophus in a higher snout-vent length, tibia length, eye-diameter, eye-to-
nostril distance, Toe III length, and Toe IV length (Table 4). The average eye-diameter, tympanum diameter, Toe III
length, Toe IV length was significantly higher in P. stictus in comparison with P. uranobates, but eye-to-nostril
distance was significantly lower in P. stictus compared to P. uranobates (Table 4).
TABLE 4. Mean and standard deviation of measurements showing with significant differences in Tukey Test HSD.
Remarks. Variation in vomerine odontophores has received some attention in the literature (e.g., Lynch, 1980;
Lynch and Duellman, 1997; Lynch, 2001; Duellman and Lehr, 2009). Lynch and Duellman (1997) compared the
vomerine odontophores among several genera of Brachycephaloidea and discussed the loss of this character in
some taxa, as in Atopophrynus, Geobatrachus, Noblella, Bryophryne, and a few species in the genus of Phrynopus,
Psychrophrynella, Eleutherodactylus and Craugastor (Figure 8A–C) (Lynch, 1975; Lynch, 2001; Hedges et al.
2008; Trueb and Lehr, 2008; Lehr and Catenazzi, 2008 and 2009; Padial et al. 2014a; Catenazzi et al. 2015). Lynch
and Duellman (1997) also noted that species of Pristimantis from western Ecuador possess small odontophores,
Variables Wilks'
Lambda
Partial
Lambda
F-remove (2.39) P Root 1 Root 2
Snout vent length 0.033 0.773 5.278 0.00 1.2387 -0.0678
Tibia length 0.034 0.752 5.929 0.00 0.8635 -0.3502
Eye diameter 0.029 0.882 2.403 0.10 0.3053 -0.4290
Eye to nostril distance 0.035 0.718 7.034 0.00 1.0696 0.0121
Snout eye distance 0.040 0.635 10.309 0.00 -1.3239 0.2792
Tympanum diameter 0.036 0.705 7.496 0.00 -0.9808 0.0280
Toe III length 0.035 0.723 6.867 0.00 -1.2393 -0.9580
Toe IV length 0.030 0.840 3.424 0.04 1.2585 0.7482
Eig 10.1724 2.4713
Cp 0.8045 1
Variables P. uranobates P. leptolophus
Mean SD P Mean SD P
Snout vent length 24.9 1.9 0.65 23.4 1.4 0.00
Tibia length 12.5 0.8 0.26 11.3 0.5 0.00
Eye diameter 2.9 0.2 0.00 2.7 0.1 0.00
Eye to nostril distance 2.5 0.2 0.00 2.1 0.1 0.00
Snout eye distance 4.0 0.3 0.99 4.0 0.2 0.95
Tympanum diameter 1.2 0.1 0.03 1.3 0.2 0.50
Toe III length 3.9 0.3 0.00 3.8 0.3 0.00
Toe IV length 6.5 0.5 0.00 6.3 0.4 0.00
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which are concealed beneath the palate tissue—a very distinctive condition for the genus. But despite the variation
and potential importance of these features in the taxonomy of brachycephaloids (and in other taxa as well),
differences in terminology have obscured the interpretation of observed variation. For example, Lynch (e.g., Lynch,
1979, Lynch, 1980; Lynch, 1981) equates the term dentigerous process of the vomers with vomerine odontophores
in many of the descriptions of brachycephaloids frogs. Lynch and Duellman (1997) also treated vomerine
odontophores as a synonym of the dentigerous process of the vomer, although, they recognized that when poorly
discernible, the odontophore could be present but concealed beneath the mucosa. Lynch (2001) makes some
distinctions between odontophore and dentigerous process. Nevertheless, none of them provided accurate
definitions. Only Savage (2002) defined the odontophore as tooth-bearing processes of the vomer. Duellman et al.
(2006) and Duellman & Lehr (2009) reviewed the terminology and proposed using dentigerous process to replace
vomerine odontophores. As a result of the inconsistence in the use of terminology, some species have been
described as lacking dentigerous process when the process was in fact concealed underneath the mucosa (Figure
8A–C) (see also Duellman and Hedges, 2007; Lehr, 2007).
FIGURE 7. Graphic representation of the two roots of the discriminant analysis for morphometrics.
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FIGURE 8. Vomers variation of some brachyphaloids frogs. A. Eleutherodactylus nitidus (KU-102657), lacks the dentigerous
process; B. Eleutherodactylus leprus (KU-55963), dentigerous process represented by long sheet, the odontophores are absent;
C. Craugastor pygmaeus (KU-86876), dentigerous process represented by sharp and elongated structure, the odontophores are
absent; D. Craugastor escoses, adult female (KU-117355), dentigerous process consists mainly of a large odontophore; E.
Pristimantis danae, adult male (KU-164062; SVL: 36 mm), elongate dentigerous process with small odontophore; F.
Pristimantis orpacobates, adult female (KU-170134; SVL: 43 mm), dentigerous process with midsize odontophore; G.
Pristimantis orestes, adult female (KU-141995; SVL: 23.5 mm), elongate dentigerous process with low odontophore; H.
Pristimantis vicarius, adult male (KU-170155; SVL: 34 mm), very elongated dentigerous process with odontophore in the
distal end. Bars represent measurements of 2 mm; gray denotes cartilage and stippling denotes bones. (All drawings based on
original illustrations by Lynch).
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Another important osteological feature in the taxonomy of brachycephaloid frogs are the oblique keels. Lynch
(1980) first mentioned oblique keels but did not define this character. I define the oblique keels as a ridge
concealed or not by the buccal mucosa, and formed by the anterior elongated portion of the posterior process of
vomers (Figure 5). It is important to note that the oblique keels and the vomerine odontophores are not
homologous. Oblique keels are exposed anteriorly to the odontophores in the posterior process of vomers, while
odontophores and teeth may be present or not.
The vomerine odontophore is the ossified and exposed part of the posterior vomer processes. It may bear teeth
and its structure is variable. In brachycephaloids, the dentigerous process can be either absent (Figure 8A—some
Eleutherodactylus and Noblella), short and mainly formed by the odontophores (Figure 8D), elongated and of
different size, shape, and position, but bearing the odontophore at the posterior end (Figure 8E–H), or lacking the
odontophore (Figure B–C) (e.g., Bryophryne cophites, Craugastor pygmaeus, Eleutherodactylus marnockii, E.
rufescens, E. longipes, E. dixoni, E. leprus). Pristimantis of the P. leptolophus group of the paramos, such as P.
stictus, P. leptolophus, and P. lasalleorum, lack odontophores and vomerine teeth. A thorough study of the
variation of these characters in brachycephaloid frogs is required, especially in the poorly know genus Pristimantis.
On the other hand, not only is the morphology of Pristimantis poorly known, but also its species diversity and
geographic distribution. For example, the paramos of Colombia have so far been little explored, and the knowledge
on amphibian communities in the different highland areas is rudimentary. The allopatric distributions of species on
paramo areas of mountain tops suggest an important role of vicariant speciation in highland species. Indeed, the
presence of endemic species (Figure 6) (Bernal & Lynch 2008) in isolated paramos suggests that other undescribed
species could await discovery in the many isolated and unexplored paramos of the Cordillera Central of Colombia.
Acknowledgements
For cooperation in the field, I am grateful to Fernando Castellanos, Jonathan Castellanos, David Vallejo, Julian
Rojas, Sergio Escobar and Airan Arango. To Aguas de Manizales and Jorge Uribe for cooperation in the field
work. To Diego Cisneros Heredia, Mariane Targino, Pedro Diaz Dos Santos, Marco Rada and José Padial for their
feedback and comments. To Professor John D. Lynch for comments and allowing access to the amphibians
laboratory in the Instituto de Ciencias Naturales. To Rafe Brown for providing a photo collection of the
Biodiversity Institute at the University of Kansas. To Alan Resetar for providing photos of types specimens from
the Field Museum of Natural History. Prof. Lynch also permitted me to have free access to his largely unpublished
drawings of brachycephaloids skulls.
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APPENDIX 1. Specimens examined. Abbreviations: holotype (H); paratype (P); male (M); female (F).
Pristimantis lasalleorum: COLOMBIA: ANTIOQUIA: Frontino: FMNH-69719-20 (P–F).
Pristimantis leptolophus: COLOMBIA: CAUCA: Belalcazar: 41857-58 (F), 41865 (F), 41898 (F); Coconuco: 07799 (F); Inza:
11487 (F), 41831 (F), 41834-35 (F), 41838-40 (F); Paez: ICN-06745 (F), 06750-51 (F), 06753 (F), 06762 (F), 06765 (F),
07033 (F), 7058 (F), 07304-05 (F); Purace: KU-169041 (H–F).
Pristimantis maculosus: COLOMBIA: ANTIOQUIA: Belmira, ICN-8591 (H–F), ICN-8592-93 (P–M); Sonson: ICN-8594(P–
F), 8596 (P–M); CALDAS: Manizales: ICN-55578 (F).
Pristimantis parectatus: COLOMBIA: ANTIOQUIA: Bello: ICN-9247 (P–F); Sonson: ICN-9248 (H–F), ICN-55743 (F);
CALDAS: Aguadas: ICN-55740 (F); Pensilvania: ICN-41687-90 (P–F), ICN-41691-95 (P–M).
Pristimantis peraticus: COLOMBIA: VALLE DEL CAUCA: Tenerife: KU-168915 (H–F); Tulua: ICN-40772-73(F), ICN-
40777 (F); TOLIMA: Rio Blanco: ICN-55742 (F).
Pristimantis scoloblepharus: COLOMBIA: ANTIOQUIA: Belmira: ICN-8583 (H–F), ICN-8584-85 (M–P); Sonson: ICN-
18770-71 (M–P), ICN-18774 (M–P), ICN-55744 (F).
Pristimantis uranobates: COLOMBIA: CALDAS: Manizales: 55580-81 (F), 55587-88 (F); Villamaria, ICN-14424 (H–F), ICN
14425 (P–M), 14426-27 (P–F) 14428-31 (P–M); QUINDIO: Salento: ICN-24974-24977 (F), 24980 (F), 24983-84 (F),
29836-38 (F), 29843-46 (F), 29873 (F), 29875-76 (F), 29881 (F).
