A new small frog species of the genus Pristimantis (Anura: Craugastoridae) from the northern paramos of Colombia Author González-Durán, Gustavo A. text Zootaxa 2016 4066 4 421 437 journal article 38713 10.11646/zootaxa.4066.4.4 d7e1540e-9a08-41e6-8d31-408b77fdf9c1 1175-5326 258317 03DC485E-A9B6-4E5B-93D8-A7639BB4BF22 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, 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 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 s tictus 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). 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. TABLE 1. Diagnostic morphological characters in the Pristimantis leptolophus species group.

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
continued.

Lateralfringes on finger	Nuptial pads on thumb	Ulnar tubercles	Tarsal tubercles	Heel tubercle	Tubercles on upper eyelid
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
continued.

view

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	Present	Absent	Obliques	Lynch, 1991 , this work

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 ( x = 19.9 ± 1.2; n=15) and females 22.0–28.2 ( x = 25.4 ± 1.6; n=17). 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 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 5 C–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). Variation. 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 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

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 . 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; X2 = 170.09; df= 44 p <0.00) and second root (R= 0.85; X2 = 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).

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 Pristimantis stictus differs from P . leptolophus in a higher snout-vent length, tibia length, eye-diameter, eye-tonostril 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. uranobates leptolophus

Variables	P.	P.
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

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 8 A–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, 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 8 A–C) (see also Duellman and Hedges, 2007; Lehr, 2007 ). FIGURE 7. Graphic representation of the two roots of the discriminant analysis for morphometrics. 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). 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 8 A—some Eleutherodactylus and Noblella ), short and mainly formed by the odontophores ( Figure 8 D), elongated and of different size, shape, and position, but bearing the odontophore at the posterior end ( Figure 8 E–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 .