Osteocephalus vasquezi, Venegas & García-Ayachi & Toral & Malqui & Ron, 2023

Venegas, Pablo J., Garcia-Ayachi, Luis A., Toral, Eduardo, Malqui, Jose & Ron, Santiago R., 2023, A new species of spiny-backed tree frog, genus Osteocephalus (Anura, Hylidae), from the Yanachaga Chemillen National Park in central Peru, Evolutionary Systematics 7 (2), pp. 237-251 : 237

Osteocephalus vasquezi sp. nov.

Figs 2 , 4 , 5 , 6A-D

Type material.

Holotype. Peru • Adult male; Pasco department, Oxapampa province, National Park Yanachaga Chemillén, Quebrada Honda (close to the park rangers’ checkpoint of Huampal); 9°48'53"S, 75°38'13"W; 1000 m; 15 Aug. 2010; P.J. Venegas, V. Duran, and L. Lujan leg.; CORBIDI 7284 (Fig. 2A, B ).

Paratypes. Peru • 5 ♂♂ adults, 2 ♀♀ adults, 2 juveniles, collected with the holotype; CORBIDI 7270, 7272, 7283, 7285-86, 7271, 7280, 7304, 7438 • 3 ♂ adults, 1 ♀ subadult, collected at the same location of the holotype; 16 Aug. 2010; P.J. Venegas, V. Duran, and L. Lujan leg.; CORBIDI 7277-79, 7275 • 2 ♂♂ adults; Pasco department, Oxapampa province, Quebrada Shuler; 10°10'22"S, 75°34'13"W; 1050 m; 18 Aug. 2010; P.J. Venegas, L. Lujan, and C. Landauro leg.; CORBIDI 7273-74 • 2 ♂♂ adults; Pasco department, Oxapampa province, Quebrada Gallito; 10°13'27"S, 75°35'3"W; 1010 m; 20 Aug. 2010; P.J. Venegas, V. Duran, and C. Landauro leg.; CORBIDI 7276, 7282 • 1 ♂ adult; Pasco department, Oxapampa province, Trocha Colonos; 10°11'03"S, 75°34'27"W; 1050 m; 27 Aug. 2010; P.J. Venegas, V. Duran, L. Lujan, and C. Landauro leg.; CORBIDI 7281.

Definition.

Osteocephalus vasquezi sp. nov. can be distinguished from its congeners by the following combination of characters: (1) size sexually dimorphic; maximum SVL in males 52.9 mm (n = 14), in females 75.5 mm (n = 2); (2) skin on dorsum of breeding males bearing conical tubercles with keratinized tips, not present in non-breeding males, smooth in females; (3) skin on flanks weakly areolate in the anterior two-thirds and smooth posteriorly; (4) hand webbing formula varies from II2--3-III 2½ -2+IV to I basal II2---3+III3- 2½ IV; foot webbing formula varies from I1--1II1--1-III0-1-IV1--1-V to I1-2-II1-2-III2--2-IV2---1V; (5) in life, dorsum varies from brown to dark brown or orange-brown, with or without dark brown irregular marks; (6) throat brown or tan with a distinctive pattern of white irregular blotches or vermiculations, as well as tan to brown blotches on a whitish cream to creamy tan background; chest and belly cream or creamy tan with chocolate blotches or flecks; (7) cream suborbital mark indistinct, clear labial stripe distinct or faint; (8) color of dorsolateral region of flanks similar to dorsal coloration; ventrolateral region whitish cream or brownish cream with brown scattered blotches and/or vermiculations; (9) dermal roofing bones of the skull not exposed; (10) in life, bones green; (11) in life, iris dark brown with golden vermiculations or flecks; (12) vocal sacs paired, small, located laterally, behind jaw articulation; (13) in life, juveniles with red iris, dorsal surface of body and limbs dark brown (almost black) with marks or coppery with dark brown marks, without conspicuous pale elbows, knees, and heels; (14) larvae with LTRF of 3/9.

Diagnosis.

Osteocephalus vasquezi sp. nov. is most similar to O. mimeticus and O. aff. mimeticus (see Ortiz et al. 2022). Both species are similar in having dark irises with golden marks (see Fig. 2A-F , 3E-H ). However, the new species can be distinguished from O. mimeticus in having a cream or creamy-tan venter with a well-defined pattern of brown chocolate blotches and flecks (venter cream, tan, or brown without marks in O. mimeticus ). Additionally, adult males of Osteocephalus mimeticus are larger than those of O. vasquezi sp. nov. with non-overlapping ranges [SVL in O. mimeticus 58.2-67.5 mm, mean = 63.4 (n = 7), versus 40.9-52.9 mm, mean = 47.9 (n = 13)]. Furthermore, the tadpoles of O. vasquezi are strikingly different from the tadpoles of O. mimeticus (see Fig. 6 ) (state of characters between parenthesis) by having: truncate snout in dorsal view (rounded), nostrils oriented dorsolaterally closer to the eyes (nostrils oriented laterally and closer to the end of snout), fins at the posterior third of tail conspicuously narrower (gently narrower), and a higher number of lower labial tooth rows LTRF = 3(3)/9 (LTRF = 2(2)/6).

Other species similar to Osteocephalus vasquezi sp. nov. are O. festae and O. verruciger (Fig. 3 ), both species of the Osteocephalus buckleyi group have predominantly dark irises, tuberculate dorsal skin, and brown dorsal coloration. Although some O. festae and O. verruciger share with O. vasquezi sp. nov. a cream or brownish-cream venter with dark brown chocolate marks, they differ from O. vasquezi sp. nov. in having a dark brown iris without bright marks and areolate flanks (iris dark brown with golden vermiculations or flecks, and smooth to weakly areolate in the anterior third of the flank in O. vasquezi sp. nov.). Additionally, in O. festae and O. mimeticus the subocular mark is usually conspicuous, while in the new species it is faint.

Osteocephalus mutabor , O. omega , and O. sangay Chasiluisa, Caminer, Varela-Jaramillo & Ron, 2020, from the O. buckleyi group, also have brown dorsal coloration. However, in life, irises of O. mutabor and O. sangay are bronze with irregular black reticulations and in O. omega are golden yellow, whereas in the new species the irises are dark brown with golden vermiculations or flecks. Osteocephalus mutabor also can be readily distinguished by having a dorsal pattern of distinctive transversal stripes (absent in O. vasquezi sp. nov.) and lacking dark blotches or flecks on the ventral surface. Additionally, metamorphs and juveniles of O. mutabor have green dorsal coloration (black with golden marks in O. vasquezi sp. nov.). Females of O. sangay have scattered tubercles on dorsum, while females of O. vasquezi sp. nov. have the skin of dorsum smooth. Although O. sangay also has cream or tan venter with dark brown dots, the new species can be distinguished by its distinctive pattern of white or brown irregular blotches or vermiculations on the throat.

The ventral coloration of Osteocephalus vasquezi sp. nov. is shared by some individuals of O. buckleyi Boulenger, 1882, O. cabrerai , O. camufatus , O. cannatellai Ron, Venegas, Toral, Read, Ortiz & Manzano, 2012, Osteocephalus duellmani , O. germani Ron, Venegas, Toral, Read, Ortiz & Manzano, 2012, O. helenae Ruthven, 1919, and O. vilmae Ron, Venegas, Toral, Read, Ortiz & Manzano, 2012. However, the new species can be easily distinguished, in life, in having a dark brown iris with golden vermiculations or flecks (iris varies from cream to golden or reddish golden and yellow with or without irregular reticulations in the afore listed species, except by O. duellmani due to its coloration in life is unknown). Furthermore, O. buckleyi , O. cabrerai , O. camufatus , O. cannatellai , O. helenae , and O. vilmae differ from O. vasquezi sp. nov. in having prominent tarsal tubercles (indistinct or absent in the new species). Osteocephalus cabrerai also has distinct tubercles on the lower jaw and a fringe in the outer edge of Finger IV, both characters absent in O. vasquezi sp. nov. The poorly known O. duellmani and O. germani have the skin on flanks coarsely areolate or areolate, respectively, and in O. vasquezi sp. nov. only weakly areolate in the anterior third of the flank, or completely smooth. Furthermore, O. duellmani differs from O. vasquezi sp. nov. in having the dorsum shagreen (tuberculate in the new species) and a conspicuous light subocular mark (faint in O. vasquezi sp. nov.). In O. germani the throat is cream with faint brown flecks, while in the new species is cream with brown blotches or brown with cream vermiculations.

Predominantly dark brown irises are also present in O. alboguttatus Boulenger, 1882, O. heyeri Lynch, 2002, O. melanops , and O. subtilis Martins & Cardoso, 1987. Osteocephalus heyeri and O. alboguttatus can be easily distinguished from the new species by having brown flanks with scattered white blotches (flanks whitish cream or brownish cream with dark brown blotches and vermiculations in O. vasquezi sp. nov.). Osteocephalus subtilis differs from O. vasquezi sp. nov. in having the armpits, groins, anterior and posterior surfaces of thighs, and shanks blue, and the iris dark brown without bright marks. Osteocephalus melanops has a cream to white venter, while in the new species it is cream or creamy-tan with brown chocolate blotches and flecks.

Osteocephalus oophagus , and O. taurinus can be easily distinguished from O. vasquezi sp. nov. in having bronze to golden irises (clear in preservative) with black lines radiating from the pupil. Furthermore, Osteocephalus taurinus differs from O. vasquezi sp. nov. in having dermal roofing bones of the skull exposed (not exposed in the new species). Osteocephalus leprieurii , and O. yasuni Ron & Pramuk, 1999 can be distinguished by their golden to golden brown irises with fine irregular dark venations and a broad dark brown horizontal midline. Osteocephalus yasuni also can be distinguished from O. vasquezi sp. nov. by its ventral coloration (from yellow to creamy-yellow in O. yasuni vs. cream or brownish-cream with chocolate-brown blotches and flecks in O. vasquezi sp. nov.) and the color of bones (white in O. yasuni and green in O. vasquezi sp. nov.). Additionally, O. leprieurii and O. yasuni breed in ponds and flooded areas ( Jungfer et al. 2013), while the new species breeds in torrential streams.

Osteocephalus castaneicola Moravec, Aparicio, Guerrero-Reinhard, Calderón, Jungfer & Gvoždík, 2009, O. deridens Jungfer, Ron, Seipp & Almendáriz, 2000, O. fuscifacies Jungfer, Ron, Seipp & Almendáriz, 2000, O. leoniae Jungfer & Lehr, 2001, O. planiceps Cope, 1874, and O. vilarsi differ from O. vasquezi sp. nov. (characters in parentheses) in having vocal sac single and subgular (vocal sacs paired, located above the arm and below the ear), dorsal skin not sexually dimorphic and more or less smooth in both sexes, except in O. planiceps and O. vilarsi (dorsal skin sexually dimorphic, strongly tuberculate in males and smooth in females), and breeding in phytotelmata, such as leaf axils, fruit capsules, bamboo and tree holes ( Jungfer et al. 2013; Ferrão et al. 2019) (breeding in torrential streams). Additionally, the dark brown iris with golden vermiculation or flecks in O. vasquezi sp. nov. differs from the iris of all species in the O. planiceps group. Irises are golden to bronze with fine dark reticulate or black lines radiating from the pupil in O. castaneicola , O. leoniae , O. deridens , O. fuscifacies , O. planiceps , and O. vilarsi .

Of the 29 species of Osteocephalus (Frost 2022) known to the date, only the larvae of six species have been formally described (i.e., O. cabrerai , O. festae , O. mimeticus , O. oophagus , O. taurinus and O. vilarsi ) ( Trueb and Duellman 1970; Henle 1981; Hero 1990; Schiesari et al. 1996; Ron et al. 2010; Ferrão et al. 2019). The tadpoles of O. vasquezi sp. nov. differ from its congeners by the following characters: truncate snout in dorsal view (except for O. oophagus , in the rest of species it is rounded), nostrils oriented dorsolaterally closer to the eyes than the end of snout (nostrils oriented laterally in the rest of species, closer to the end of snout in O. cabrerai , O. mimeticus , and O. vilarsi , and intermediate in O. taurinus and O. verruciger ), fins at the posterior third of tail conspicuously narrower (except for O. festae , in the other species fins are gently narrower). Furthermore, O. vasquezi sp. nov. has a higher number of lower labial tooth rows: LTRF = 3(3)/9 vs. LTRF = 2(2)/6(1) in O. cabrerai , 4-5/7 in O. festae , 2(2)/6 in O. mimeticus , 2(2)/6 in O. oophagus and 2(2)/3-7(1) in O. taurinus , 2(2)/5-6(1) in O. vilarsi , and 2(2)/5(1) in O. verruciger .

Description of holotype.

Adult male (Figs 4 , 5 ), 51.4 mm SVL, head length 15.5 mm, head width 15.5 mm, eye diameter 5.3 mm, tympanum diameter 3.1 mm, femur length 23.7 mm, tibia length 28.0 mm, foot length 22.3 mm. Head narrower than body; snout truncate in lateral and dorsal views; canthus rostralis distinct and rounded; loreal region concave; internarial area depressed; nostrils slightly protuberant, directed laterally; interorbital area flat, with scattered small tubercles, lateral margins of the frontoparietals inconspicuous through skin; eye strongly protuberant; tympanic membrane evident, slightly wider than high, separated from eye by ca. 93% of its diameter; tympanic annulus distinct. Tongue cordiform, widely attached to floor of mouth; vomerine odontophores angular, adjacent medially, posteromedial to choanae, bearing 9 and 8 (left/right) vomerine teeth; choanae with capsular shape, oblique; deflated vocal sacs distinct above the arms and below the ears.

Axillary membrane present, reaching half the arm length; ulnar tubercles absent; relative length of fingers I <II <IV <III (Fig. 4 ); fingers bearing oval discs, that of Finger III about two thirds of tympanum diameter; subarticular tubercles prominent, round to ovoid except for slightly bifid distal subarticular tubercle of Finger IV; supernumerary tubercles present, distinct; palmar tubercle elongated; prepollical tubercle protuberant, elliptical; prepollex present; dark brown, keratinous nuptial excrescences covering inner surface of prepollex up to the intercalary cartilage of thumb; webbing basal between fingers I and II; webbing formula of fingers II 2--3 III 3-- 2½ IV (Fig. 4 ).

Small tubercles on tibiotarsal articulation; dorsal surface of tarsus covered by scattered minute keratinized conical tubercles, more abundant on outer edge; small tubercles scattered along ventrolateral edge of foot; toes bearing discs slightly wider than long, smaller than those of fingers; relative length of toes I <II <V <III <IV; outer metatarsal tubercle ill defined, small, round; inner metatarsal tubercle larger, ovoid; subarticular tubercles single, round, protuberant; supernumerary tubercles restricted to the soles; webbing formula of toes I 1-1+ II 1-1+ III 1-1+ IV 1+-1 V (Fig. 4 ).

Skin on dorsum, head, and dorsal surfaces of hindlimbs shagreen, covered by conical tubercles with keratinized tips, tubercles minute on head and limbs; skin on flanks weakly areolate; skin on venter granular; skin on ventral surfaces of head smooth, on those of thighs smooth on the anterior half and granular on the posterior half, smooth on shanks. Cloacal opening directed posteriorly at upper level of thighs; short simple cloacal sheath covering cloacal opening; round tubercles around vent and on posterior surface of proximal third of thighs.

Color of holotype in life.

Based on digital photographs (Fig. 2A, B ). Dorsal surface of head dark brown, dorsum brown with dark brown irregular blotches, dorsal surfaces of limbs brown with dark brown diagonal bars; sides of head dark brown with pale lips and a thin white subocular stripe, upper flanks brown, mid- and lower-flanks dark brown with scattered vermicular white blotches. Venter light cream with chocolate blotches and flecks, more abundant on chest; throat dark brown with white vermiculations; ventral surface of limbs, palms, and soles brown. Iris dark brown with irregular golden vermiculation.

Color of holotype in preservative.

Dorsal surface of head dark brown, dorsum pale brown with dark brown irregular marks, dorsal surface of limbs pale brown with dark brown diagonal bars; sides of head dark brown with pale lips and a thin white subocular stripe, flanks dark brown with scattered vermicular white blotches. Venter dirty cream with chocolate blotches and flecks more abundant on the chest; throat brown with white vermiculations; ventral surface of limbs brownish cream, palms and soles brown.

Variation.

Variation in dorsal and ventral coloration of preserved specimens is shown in Fig. 5 . Dorsal background coloration varies from grayish-brown to brown or dark brown, irregular dark brown marks usually present. One specimen (CORBIDI 7276) has brownish-cream scattered blotches on dorsum; CORBIDI 7273 and 7278 have pale brown dorsum without dark marks; CORBIDI 7271 with scattered faint vermiculations on dorsum. Ventral surfaces of preserved specimens (Fig. 5 ) vary from light cream to dirty cream with scattered chocolate-colored blotches, spots, or flecks; one specimen (CORBIDI 7274) has a cream venter with faint blotches. Some specimens (e.g., CORBIDI 7275, 7277) have venters densely spotted, the two adult females (CORBIDI 7271, 7280) have finely reticulated venters. Throat coloration light cream with faint chocolate blotches on sides of throat (CORBIDI 7271, 7282), or light cream with scattered chocolate blotches (CORBIDI 7277, 7278, 7283, 7286), or dark brown (CORBIDI 7280, 7281) or pale brown with white vermiculations (CORBIDI 7276). Undersides of limbs vary from cream to brownish-cream. Cream-colored low tubercles on the outside edge of the forearm present only in CORBIDI 7275. The skin of the anterior and posterior surfaces of thighs and concealed surfaces of shanks is light cream or brownish-cream. The vent region is usually dark brown surrounded by a light cream edge; CORBIDI 7283 has the vent region dirty cream. Background color of flanks varies from grayish-cream to brownish-cream; some specimens (CORBIDI 7273, 7282, 7280, 7271) have a long undulating longitudinal dark brown stripe along the flanks; CORBIDI 7274 has scattered circular blotches along the flanks; the rest of the specimens have irregular dark brown or pale brown blotches along the flanks. Lateral head coloration varies from brown to dark brown. The subocular white mark varies from a thin stripe (e.g., CORBIDI 7276) to a faint blotch (e.g., CORBIDI 7272), and some specimens have little brown spots on the subocular blotch (e.g., CORBIDI 7277, 7281).

The only known adult females (n = 2) lack tubercles on dorsum, while in males the dorsum varies between lacking tubercles (CORBIDI 7275, 7279) to having well defined tubercles with a keratinized spicule on the tip (e.g., CORBIDI 7283). The skin on the flanks is weakly areolate to areolate in the anterior one-third and smooth posteriorly. The two adult females (CORBIDI 7272, 7280) have smooth flanks. Head shape is truncate in dorsal and lateral view as in the holotype. The tympanic annulus is concealed dorsally by a supratympanic fold and is of a lighter color than the background. The distal subarticular tubercle on Finger IV is slightly bifid in all specimens. Measurements and proportions of the type series of Osteocephalus vasquezi sp. nov. are summarized in Table 2 . In the examined series, the largest male has a SVL of 52.9 mm and the largest female 75.5 mm; mean adult male SVL = 48.1 mm (n = 14, SD = 3.86), mean adult female SVL = 70.7 mm (n = 2).

Based on digital photographs of the adult specimens in life (Fig. 2A-F ), the dorsal background color of body and limbs varies from pale brown (CORBIDI 7273) to dark brown (CORBIDI 7272) or orange-brown (CORBIDI 7270), with or without dark brown irregular marks on dorsum and dark brown or faint brown transversal bands on limbs; dorsal surface of head is darker than body in some specimens (e.g., CORBIDI 7271), sides of head are darker than body in some specimens, especially in the tympanic region; flanks are pale brown in all specimens with the upper region brownish-cream or whitish-cream with scattered dark brown irregular blotches (ill-defined in CORBIDI 7273); throat usually whitish cream with brown or dark brown irregular blotches, some specimens (e.g., CORBIDI 7280) have the throat brown with irregular white blotches; chest and belly vary from cream to whitish-cream or dirty cream with scattered brown blotches, spots, and flecks or with fine brown reticulations on the belly (e.g., CORBIDI 7272, 7271). The nuptial pad is cream or brownish cream and contrasts with the background brown color of the thumb and rest of the fingers. Iris dark brown with golden vermiculations or flecks.

There is significant change in color between juveniles and adults. The following description is based on a digital photograph of juvenile CORBIDI 7304 (Fig. 2G ). Top of head, body, and limbs dark brown (almost black) with golden marks as dorsolateral irregular stripes, from the tip of snout to the posterior end of the sacrum, scattered vermiculations, transverse bands on limbs, and golden labial marks; venter dark brown, almost black; iris bright red.

Tadpoles.

Tadpoles were collected at Quebrada Honda, the same location as for the holotype, on 20 August 2010. These larvae belong to the exotrophic, lotic, suctorial guild as defined by Altig and McDiarmid (1999). Guild assignment was based on oral disk and body morphology. Morphometric data are provided in Table 3 .

In dorsal view, a tadpole in Stage 33 (Fig. 6A-D ) shows an elliptical body, widest between eye and spiracle, with a rounded snout. Eyes are relatively large (BL about 10.4 times larger than ED), separated by a distance 1.34 times the internarial distance, directed and positioned dorsolaterally, not visible in ventral view. External nares oval, located dorsolaterally, at about one fourth the distance between anterior margin of snout and anterior margin of eye.

In profile (Fig. 6B ) body depressed (BW/BH = 1.35), flattened ventrally, snout slightly rounded. Oral disc not emarginated. Spiracle single, sinistral, inner wall free from body, its tip closer to the vent than the eye. Spiracle opening rounded, at level of the forelimbs. Tail musculature robust up to first one third, decreasing in size towards the tip of tail. Dorsal fin does not extend onto the body, slightly convex, and reaches its highest width up the half of tail. Tail tip acuminate. Ventral fin slightly convex, beginning in the tail-body junction and tapering abruptly in the last third towards the tail tip. Vent tube medial, with both walls attached directly to ventral fin, opening directed posteroventrally. Lateral line system present on: dorsal body, middle body; supraorbital, infraorbital, posterior supraorbital; and posterior infraorbital. No glands.

Oral disc anteroventral (Fig. 6C, D ); transverse width 5.6 mm; 70.86% of BW), not emarginated, LTRF 3(3)/9; papillae distributed around oral disc; tooth rows complete except for medial gap in row A3; A1 = 778, A2 = 592, A3 = 195 + 203; P1 = 267, P2 = 325, P3 = 336, P4 = 380, P5 = 323, P6 = 409, P7 = 316, P8 = 493 and P9 = 296.

Based on PJV’s field notes of tadpoles the color in life between stages 30 to 42 (Fig. 2H , Stage 42): dorsal surface of head, body, limbs, and base of tail light coppery with black marks, flanks and sides of tail brown, venter brown and slightly translucent, iris red. Tadpoles in stages 31, 35, and 37 have a dark brown dorsum, venter brown with the gut visible; tail musculature brown, lighter than the dorsum; fins translucent with a pale brown tone. Tadpoles in stages 39 and 40 have the same coloration as of the aforementioned stages but with a pale dorsolateral stripe on each side.

Dorsum pale brown, caudal muscle beige with brown spots (Fig. 6A-C ). Eyes, flanks, spiracle, vent tube, fins, and venter dark brown surrounded by transparent areas. Venter and fin transparent.

Distribution and natural history.

Osteocephalus vasquezi sp. nov. is only known from the type locality in Cordillera de Yanachaga, Pasco department, at elevations between 1000 and 1150 m, in the upper Amazon basin of central Peru (Fig. 7 ). This new species inhabits the premontane forest of the Río Huancabamba canyon. The distribution lies within the Yanachaga Chemillén National Park and in the ecoregions of Selva Alta (400-1000 m) and Yungas (500-2300 m), according to Brack-Egg (1986) and Peñaherrera del Aguila ( 1989). All individuals were collected at night, on leaves and branches of bushes up to 2 m above the ground, along ravines in Quebrada Honda and Quebrada Shuler, both drainages of the Río Huancabamba. Sympatric amphibians were Bolitoglossa peruviana Boulenger, 1883, Leptodactylus rhodonotus Günther, 1869, Pristimantis diadematus Jiménez de la Espada, 1875, P. minutulus Duellman & Hedges, 2007, Pristimantis sp., Rhinella aff. leptoscelis Boulenger, 1912, and R. poeppigii Tschudi, 1845. Recently metamorphosed juveniles of Osteocephalus vasquezi sp. nov. were on the rocks and leaves of low bushes on the shores of Quebrada Honda on the second half of August. The ravine of Quebrada Honda has a torrential stream of clear waters and a rocky bottom. Tadpoles of O. vasquezi sp. nov. were observed adhered to rocks, presumably with their oral disk, in the narrow and very torrential portion of Quebrada Honda and in the Huancabamba River. Tadpoles of Rhinella aff. leptoscelis were found in the same stream, but in lower abundance.

Etymology.

The specific name is a patronym for Pedro Vásquez Ruesta, a Peruvian forest engineer, who is a pioneer in the wildlife management in Peru. Since 1978 he has worked for the development of wildlife management and protected natural areas as a professor at the Faculty of Forestry at Universidad Nacional Agraria La Molina, Lima, Peru, teaching to generations of forest engineers about wildlife management and conservation. During his academic life, Pedro Vásquez Ruesta made many contributions to the field of conservation of natural resources, advising theses and published scientific articles especially about the management of caimans and deer.

Figures

Figure 2. Dorsolateral and ventral views of Osteocephalus vasquezi sp. nov. A, B. Dorsolateral and ventral view of holotype CORBIDI 7284, SVL = 51.4 mm; C, D. Dorsolateral and ventral view of adult male, CORBIDI 7285, SVL = 40.8 mm; E, F. Dorsolateral and ventral view of adult female CORBIDI 7280, SVL = 65.9 mm; G. Dorsolateral view of a recently metamorphosed CORBIDI 7304, SVL = 13 mm; H. Dorsolateral view of tadpole in stage 42. Photographs by P. J. Venegas.

Figure 3. Dorsolateral and ventral views of adult specimens of the four most similar species to Osteocephalus vasquezi sp. nov. A, B. Male of O. festae CORBIDI 758; C, D. Female of O. festae CORBIDI 761; E, F. Male of O. mimeticus CORBIDI 9972 from Mishquillaquillo, San Martin; G, H. Female of O. mimeticus CORBIDI 9970; I, J. Male of O. verruciger CORBIDI 9477; K, L. Female of O. verruciger CORBIDI 9525 from Cordillera de Kampankis, Loreto. Photographs A-H by P. J. Venegas and I-L by A. Catenazzi.

Figure 4. Ventral views of right hand and foot of Osteocephalus vasquezi sp. nov. holotype (CORBIDI 7284). Scale bar: 10 mm.

Figure 5. Preserved specimens of Osteocephalus vasquezi sp. nov. showing variation in dorsal and ventral pattern. Left to right, first row: CORBIDI 7271 (female), 7280 (female), 7274, 7276, and the holotype 7284 (males); second row CORBIDI 7277, 7278, 7279, 7281, 7283 (males); third row: CORBIDI 7271 (female), 7280 (female), 7274, 7276, and the holotype 7284 (males); fourth row: CORBIDI 7277, 7278, 7279, 7281, 7283 (males). All specimens are shown at the same scale.

Figure 6. Preserved tadpoles of Osteocephalus vasquezi sp. nov. and O. mimeticus. A-D. Osteocephalus vasquezi stage 33, CORBIDI 24671; E-H. O. mimeticus, stage 33, CORBIDI 24672. A, E. Dorsal view; B, F. Lateral view; C, G. Ventral view; D, H. Oral apparatus. Photographs of O. vasquezi by E. Toral and O. mimeticus of L. Garcia-Ayachi.

Figure 7. Records of Osteocephalus vasquezi sp. nov. (circles; type locality with a dot), O. festae (triangles), O. mimeticus (pentagons), O. aff. mimeticus (crosses), O. mutabor (diamonds) and O. verruciger (squares). Locality data from specimens deposited at the Centro de Ornitologia y Biodiversidad and Museo de Zoologia of Pontificia Universidad Catolica del Ecuador. (Suppl. material 1).