Pristimantis quicato, Ospina-Sarria, Jhon Jairo, Mendez-Narvaez, Javier, Burbano-Yandi, Carlos & Bolívar-G, Wilmar, 2011

Ospina-Sarria, Jhon Jairo, Mendez-Narvaez, Javier, Burbano-Yandi, Carlos & Bolívar-G, Wilmar, 2011, A new species of Pristimantis (Amphibia: Craugastoridae) with cranial crests from the Colombian Andes, Zootaxa 3111, pp. 37-48 : 38-46

Pristimantis quicato sp. nov.

( Figure 2 A, 3A, 4, 5A, 7A)

Holotype. ICN 55409, adult female, collected on 30th April 2011 by Wilmar Bolívar-G. at Reserva Natural “La Sirena” (3º 31', 76º 06' W, 2600–2900 m), Vereda la Nevera, Corregimiento de Tenjo, Municipality of Palmira, Departamento Valle del Cauca, Colombia.

Paratopotypes. ICN 55406–08, 55410–11, all adult males, collected on 31st April 2011 by Jhon Jairo Ospina- Sarria, Javier Mendez, and Carlos Burbano-Yandi.

Referred specimens. UV–CD 1917–21, Juveniles males and UV–CD 1922–24, subadults males, collected with the type series.

Generic allocation. Currently there are no morphological characters supporting the monophyly of the genus Pristimantis ( Hedges et al. 2008) . However, Pristimantis is mostly composed of small, slender, direct-developing frogs from South America, and all species resembling the new taxon have been placed in the genus Pristimantis . Thus, despite we assume that additional phylogenetic evidence might eventually lead to changes in the generic allocation of the new species, assigning it to Pristimantis is the most prudent and conservative option.

Diagnosis. Pristimantis quicato sp. nov. presents the following characteristics: (1) dorsal skin granular with some conical tubercles distributed uniformly, and forming ridges (H-shaped) on shoulder; ventral skin aerolate; discoidal fold present; dorsolateral folds present, fragmented into a series of tubercles from the posterolateral corner of the orbit, and reaching the anterior half of body; (2) tympanic membrane differentiated; tympanic annulus prominent, its length 1/3–1/4 eye length, with upper edge obscured by supratympanic fold; (3) snout subacuminate in dorsal view (more rounded in females), rounded in profile; canthus rostralis slightly concave; (4) upper eyelid with small round tubercles, and one large, conical tubercle; upper eyelid narrower than IOD; low cranial crests present; (5) choanae small and partly concealed by palatal shelf of maxilla; dentigerous processes of vomers oblique, positioned posterior to level of choanae and separated medially by distance less than width of the visible vomerine process, each process bearing 6–7 teeth; (6) males lacking vocal slits, but having glandular, white nuptial pads; (7) Finger I shorter than Finger II, discs on outer fingers broad, about twice the width of digit proximal to pad; (8) fingers with broad lateral fringes; outer palmar tubercle fully divided, elliptical and equal in size; round supernumerary tubercles, subarticular tubercles larger than supernumerary tubercles, tubercles round and projecting; (9) ulnar tubercles present, subconical, folds absent; (10) heel bearing small tubercles, and one large, conical tubercle; outer edge of tarsus bearing subconical tubercles; an indistinct inner tarsal fold is present on the distal quarter of tarsus; (11) inner metatarsal tubercle oval, six to seven times the size of the oval outer metatarsal tubercle; supernumerary plantar tubercles scarce, low, subarticular tubercles larger than supernumerary tubercles, of shape round and projecting; (12) toes with broad lateral fringes; webbing absent; Toe V much longer than Toe III (reaching the base of the distal subarticular tubercle of Toe IV), discs of toes slightly smaller than those of outer fingers; (13) coloration in life highly variable, dorsally brown to yellow with reddish brown markings on green background (sometimes white, see variation); venter white with dark markings, throat brown with a white inverted triangle; flanks brown to grayish brown, with black spots; two vertical brown to black bars on the upper lips, postorbital and canthal stripe dark brown; loreal region cream with green spots; interobital blotch dark; dorsal surfaces of limbs green with reddish markings; posterior surfaces of thighs white with dark bars; anterior surfaces of thighs and groin cream with dark spots; iris reddish orange with black reticulum; (14) SVL in males 23.1–30.1 mm (mean = 26.4 ± 2.5, n = 5), in a single female 37.3 mm.

Pristimantis quicato is similar to P. calcaratus , P. kelephus and P. jubatus ( Figure 2 , 3 , 5 , 7 ). It differs from those species by a combination of traits that are described as follows (characters in parenthesis for P. quicato ). Pristimantis calcaratus ( Figure 1 , 2 B, 3B, 5B, 7D) has short vocal slits in males (absent), lacks cranial crests (present), and is smaller: snout-vent length 19.0– 23.8 mm in adult males (n=18), 27.7–31.2 mm in females (n=3) (23.1–30.1 mm in adult males and 37.3 mm in one female). Pristimantis kelephus has the dorsal skin with many round pustules forming a distinct pattern above shoulders and transverse ridges above sacrum (the tubercles are distributed uniformly across the dorsum and forming ridges only on shoulder) ( Figure 2 A, 2C); the venter is mostly black or brown with white spots (venter white with brown markings), and it lacks pale spots or bars on the posterior surfaces of the thighs (posterior surfaces of thighs white with dark bars) ( Figure 2 A, 2C). Also, P. kelephus has vocal slits (absent), the snout shape is round in dorsal view (subacuminate in males) ( Figure 2 C, 3C, 5D, 7C), the distal tip of Toe V reaches between the penultimate and distal subarticular tubercle of Toe IV (the distal end of Toe V reaches the distal subarticular tubercle of Toe IV) ( Figure 3 C), and is smaller in size: snout-vent length 18.8–21.9 mm in adult males (n=22), 27.7–30.2 mm in females (n=16) (23.1–30.1 mm in adult males and 37.3 mm in one female). Pristimantis jubatus has pustules above the sacrum, while the rest of the dorsum is smooth, and has subconical tubercles scattered on head and close tympanum (tubercles are uniformly distributed across the dorsum and form ridges only on shoulder) ( Figure 2 A, 2D); it lacks sub-anal tubercles and a conical interocular tubercle (both are present) ( Figure 2 A, 2D, 5A, 5C); the snout shape is round in both dorsal and lateral profiles (snout subacuminate in dorsal view) ( Figure 2 D, 3D, 5C, 7B); the distal tip of Toe V reaches the point between the penultimate and distal subarticular tubercle on Toe IV (the distal end of Toe V reaches the distal subarticular tubercles of Toe IV) ( Figure 3 D); and adult males and females are slightly smaller in snout-vent length, 19.8–24.1 mm in adult males, 32.4–34.6 mm in females (SVL in adult males 23.1–30.1 mm and in a single female 37.3 mm.).

Description of the holotype ( Figure 2 A, 3A, 7A). An adult female, 37.3 mm SVL. Head as wide as body, head width 41% of SVL; slightly wider than long, head length 35.9% of SVL. Snout slightly longer, rounded in dorsal and lateral view; canthus rostralis slightly concave; loreal region concave with small tubercles; lips not flared; nostrils protuberant, directed laterally; internarial distance 72.3% of eye-to-nostril distance; eye-to-nostril distance 102.1% of eye-length. Upper eyelid width slightly narrower than IOD, 50.9% of IOD; upper eyelid with small tubercles, and one large, conical tubercle on each eyelid, and low cranial crest. Tympanic annulus evident, its length 41.3% of eye diameter; supratympanic fold originating at corner of eye, encroaching on the top edge of the tympanic membrane; postrictal tubercles conical. Choanae small, oval and partly concealed by palatal shelf of maxilla; dentigerous processes of vomers slightly larger than choanae, oblique, positioned posterior to level of choanae and separated medially by distance less than width of odontophore, each process bearing 7 teeth; tongue longer than wide, posterior border notched, posterior 2/3 free. Skin on dorsum granular with some conical tubercles distributed uniformly, and forming ridges (H-shaped) on shoulders; dorsolateral folds on anterior half of body; ventral skin areolate, discoidal fold evident, throat like venter but with a diffuse black inverted triangle; skin on flanks areolate; cloacal sheath and tubercles present; ulnar tubercles present, not forming a distinct fold; thenar tubercle oval, smaller than divided palmar tubercles; supernumerary palmar tubercles present; subarticular tubercles prominent; fingers bearing lateral keels; Finger I shorter than Finger II; tips of digits round to subtruncated; inner digits bearing much narrower discs; all fingers bearing pads on digital tips.

Hind limbs moderate, heels overlap when held at right angles to sagittal plane; tibia long 59.5% of SVL; foot long 58% of SVL. Relative length of adpressed toes IV> V> III> II> I; tip of Toe V extending to base of distal subarticular tubercle of Toe IV; tip of Toe III extending to distal border of penultimate subarticular tubercle on Toe IV; discs of toes slightly smaller to those on outer fingers, bearing broad lateral fringes; webbing absent. Heel bearing small tubercles, and one large, conical tubercle; subconical tubercles along outer border of tarsus, inner edge of tarsus bearing a tubercle on distal third of tarsus; indistinct inner tarsal fold on distal 1/4 of tarsus; inner metatarsal tubercle twice as long as wide (six times larger than the subconical outer metatarsal tubercle); few supernumerary plantar tubercles present, at bases of toes II–V; protuberant subarticular tubercles.

Variation. Males are smaller than females; measurements and proportions are presented in Table 1 . The skin on the dorsum and upper surfaces of the limbs have conical tubercles, which are usually smaller on the limbs than on the dorsum of the body in females, and are uniformly distributed in males. Juveniles differ from conspecific adults by having more tubercles, but together with adult present pustules in the skin on flanks and posterior surfaces of shanks. Cranial crest with dermal ridge (ICN 55406, ICN 55408, Figure 4 A, 4C) and without dermal ridge (ICN 55407, ICN 55410–11, Figure 4 B, 4D, 4E). The coloration in Pristimantis quicato is highly variable, presenting various shades of brown or gray dorsally with diffuse dark marks, which consist of spots and longitudinal stripes (ICN 55407–08). This species has also two patterns of polymorphisms according to the scheme proposed by Carrillo-Rivera (2006) for Eleutherodactylus : (1) “Head Cap” (ICN 55406), which describes a dark area that covers the head and ends up forming a A or a inverted U shape on the back, the rest of the body white with brown spots ( Figure 4 F) and (2) Dorsoconcolor + Vertebral Dark Line (ICN 55411), where the dorsal surface of the snout, eyelids, head, and dorsum, varies from cream to brown tones differing from the rest of the darker body ( Figure 4 G). Pristimantis quicato presents variation in size and pigmentation of testis: two adult males had testis of equal size to kidneys (ICN 55407, ICN 055411), and three had size approximate to 2/3 of kidneys (ICN 55406, ICN 55408, ICN 55410); for juveniles and subadults, testis size was ranged between 1/3–1/2 of that of kidneys (UV–CD 1917–24); only one adult male was found with melanophores in a testicle (ICN 55410).

Coloration of the holotype. In life, dorsal color is light brown to green with dark markings; canthal and postorbital stripe black; interorbital blotch black; well-defined dark brown labial bars from eye to upper lip; posterior surfaces of thighs dark brown with black markings. Belly, anterior surfaces of thighs and concealed surfaces of tibia with black mottling on white background; throat brown with a white inverted triangle; flanks grayish brown, with black spots. Palms and soles grayish brown and iris reddish orange. In preservative, dorsal surfaces are grayish brown with darker spots, dark brown around dorsolateral folds; brown canthal stripe; dark labial bars; black supratympanic stripe; black cloacal patch; limbs brown with dark oblique bars; posterior surfaces of thighs grayish brown, groin, anterior surfaces of thighs white with brown blotches. Belly white with dark markings; throat brown with an inverted white triangle.

Distribution and natural history. The species is only known from the type locality ( Figure 6 ). Specimens were found between 1900 h and 2300 h in primary cloud forest in association with plants along streams, at heights above 1.5 m. The new species was not found in zones of habitat intervention (pastures, edges of road), which suggests little or no tolerance to habitat disturbance. Pristimantis quicato was found in sympatry with Pristimantis boulengeri , P. piceus , P. thectopternus , P. uranobates , Hyloscirtus larinopygion , Osornophryne sp. and Bolitoglossa sp.

Etymology. The specific epithet is used as a noun in apposition. The name quicato is commonly used by local from the departments of Cauca and Valle del Cauca to refer to children.

Figures

FIGURE 1. (A): Holotype of Pristimantis calcaratus (Boulenger 1908), adult male, BMNH 1947.2. 15.53, SVL 17.6 mm. (B): Dorsolateral and ventral view, and drawings of snout shape in dorsal and lateral view (original drawings by John D. Lynch).

FIGURE 2. Dorsolateral and ventral view of the species mentioned in the text. (A): Preserved holotype of Pristimantis quicato sp. nov. (ICN 55409, Adult female, SVL 37.0 mm). (B): Adult female of Pristimantis calcaratus (ICN 36940, SVL 27.7 mm). (C): Holotype of Pristimantis kelephus (ICN 39637, Adult female, SVL 30.2 mm). (D): Holotype of Pristimantis jubatus (ICN 52478, Adult male, SVL 22.1 mm).

FIGURE 3. Relationship in length Toe IV vs Toe V. (A): Holotype of Pristimantis quicato sp. nov. (ICN 55409, SVL 37 mm). (B): Pristimantis calcaratus (ICN 36940, SVL 27.7 mm). (C): Holotype of Pristimantis kelephus (ICN 39637, SVL 30.2 mm). (D): Holotype of Pristimantis jubatus (ICN 52478, SVL 22.1 mm).

FIGURE 4. Pristimantis quicato sp. nov. Intrapopulation variation in dorsal coloration in preservative (all adult males). (A): ICN 55406, SVL 26.0 mm. (B): ICN 55411, SVL 23.1 mm. (C): ICN 55408, SVL 26.7 mm. (D) ICN 55410, SVL 30.1 mm. (E) ICN 55407, SVL 26.1 mm. (F-G): Polymorphic patterns, “ Head Cap ” (ICN 55406) and Dorsoconcolor + Vertebral Dark Line (ICN 55411).

FIGURE 5. (A): Pristimantis quicato sp. nov. (ICN 55410, paratype, adult male, SVL 30.1 mm); (B): Pristimantis calcaratus (not collected, Reserva Forestal Bitaco, Valle del Cauca); (C): Pristimantis jubatus (not collected, PNN Munchique, Cauca, Photo: J. C. García); (D): Pristimantis kelephus (not collected, Serranía de los Paraguas, Valle del Cauca, Photo: Wilmar Bolívar-G).

FIGURE 6. Map of Colombia showing distribution of species mentioned in the text, and with an square signaling the type locality of Pristimantis quicato sp. nov.

FIGURE 7. Ornamentation of frontoparietals in: (A) Pristimantis quicato sp. nov. (ICN 55409, SVL 37 mm); (B) P. jubatus (ICN 52482, SVL 33.3 mm) and (C) P. kelephus (ICN 39639, SVL 30 mm). (D) Ornamentation of frontoparietals is absent in P. calcaratus (ICN 39640, SVL 27.2 mm).

Tables

TABLE 1. Morphometrics and proportions (in percentages) of Pristimantis quicato sp. nov. Mean and standard deviation in parentheses follow ranges.

	Female	Males
SVL	37.3	23.1–30.1 (26.4 ± 2.5)
HW	15.5	10–12.8 (11.2 ± 1.0)
HL	13.4	8.6–10.4 (9.5 ± 0.7)
IOD	7.7	5.2–6.4 (5.5 ± 0.5)
UEW	3.2	2.4–2.9 (2.6 ± 0.2)
ED	4.6	2.7–3.9 (3.3 ± 0.5)
END	4.7	2.5–3.9 (3.4 ± 0.5)
IND	3.4	2.1–3.2 (2.7 ± 0.4)
TD	1.9	1.0–1.5 (1.2 ± 0.2)
TL	22.2	15.0–17.7 (16.1 ± 1.0)
FL	22	13.4–18.3 (15.2 ± 1.8)
Head width/head length	114	113.1–123.1
Head width/SVL	41	41.5–43.4
Eye-to-nostril distance/HW	30	25–32.7
Interorbital distance /HW	50	47.3–53
Internarial distance /IOD	44	39.6–53.8
Tympanum diameter /ED	41	33.3–38.7
Tibia length/SVL	59	58.8–64.9
Foot length/SVL	59	57.3–60.8