Osteocephalus germani, Ron, Santiago R., Venegas, Pablo J., Toral, Eduardo, Morley Read,, Diego A. Ortiz, & Manzano, Andrea L., 2012

Ron, Santiago R., Venegas, Pablo J., Toral, Eduardo, Morley Read,, Diego A. Ortiz, & Manzano, Andrea L., 2012, Systematics of the Osteocephalusbuckleyi species complex (Anura, Hylidae) from Ecuador and Peru, ZooKeys 229, pp. 1-52 : 18-22

Osteocephalus germani sp. n.

Holotype.

(Fig. 13, 14) CORBIDI 05462, adult male from Peru, Región Cusco, Provincia La Convención, near Pongo de Mainique in the vicinity of Santuario Natural Megantoni (12.2581°S, 72.8425°W), 670 m above sea level, collected by G. Chavez on 23 April 2010.

Paratopotypes.

(Fig. 15 A, C) CORBIDI 06633, adult female, and CORBIDI 06660, adult male, collected with the holotype; CORBIDI 05505, adult female, collected by G. Chavez on 8 November 2009.

Paratypes.

(Fig. 15 B, D) Peru: Provincia La Convención, Comunidad Nativa Poyentimari (12.18853°S, 73.00092°W), 725 m above sea level, CORBIDI 08267, 08284, adult females, collected by G. Chavez and D. Vasquez on 28 November 2010; Comunidad Nativa Chokoriari (11.9569°S, 72.9410°W), 434 m above sea level, CORBIDI 08059, adult female, collected by D. Vasquez on 8 December 2010.

Diagnosis.

Throughout this section, coloration refers to preserved specimens unless otherwise noted. Osteocephalus germani is a medium-sized species of Osteocephalus having the following combination of characters: (1) size sexually dimorphic; maximum SVL in males 41.45 mm (n = 2), in females 50.76 (n = 2); (2) skin on dorsum bearing tubercles in males, smooth in females; (3) skin on flanks areolate; (4) hand webbing formula varying from I basal II2 –-3–III2½ -2IV to I basal II2-3III3 –-3– IV; foot webbing formula varying from I1- 1½II1–-2III1-2IV1½-1– V to I1+-2II1+-2III1+-2+IV2-1V; (5) dorsum varying from brown with dark brown marks to ligh t gray with dark brown marks; (6) venter light cream with or without dark brown flecks; (7) cream suborbital mark present, clear labial stripe absent; (8) flanks cream to brownish cream with dark brown blotches and flecks; (9) dermal roofing bones of the skull weakly exostosed; (10) bones green in life; (11) in life, iris golden to reddish golden with fine dark reticulation; (12) paired vocal sacs, located laterally, behind jaw articulation, (13) juvenile coloration unknown; (14) larvae unknown.

Osteocephalus germani is most similar to Osteocephalus buckleyi , Osteocephalus cabrerai , Osteocephalus cannatellai sp. n., and Osteocephalus vilmae sp. n. It differs from all of them in lacking prominent tarsal tubercles (tubercles are indistinct in Osteocephalus germani ). It further differs from Osteocephalus buckleyi , Osteocephalus cannatellai , and Osteocephalus vilmae in having a white to light cream venter with or without dark brown flecks (cream with brown speckling in most Osteocephalus buckleyi and Osteocephalus vilmae ,light gray to dark brown in Osteocephalus cannatellai ). Osteocephalus germani also differs from Osteocephalus cannatellai and Osteocephalus vilmae in having more abundant and keratinized dorsal tubercles (dorsal tubercles are scattered and weakly keratinized in Osteocephalus cannatellai and Osteocephalus vilmae ) and smaller size ( Osteocephalus germani male SVL range = 41.26-41.45, n = 2; Osteocephalus cannatellai male SVL range = 38.46-57.21, n = 33; Osteocephalus vilmae male SVL range = 48.23-55.77, n = 5). Mitochondrial DNA sequences show that Osteocephalus germani is the sister species of Osteocephalus cabrerai (Fig. 1). Osteocephalus germani can be easily distinguished from Osteocephalus cabrerai by ( Osteocephalus cabrerai in parenthesis): (1) absence of prominent tubercles on the lower jaw (present), (2) smooth outer edge of Finger IV (outer edge with fringe), (3) row of inconspicuous tubercles in the outer edge of tarsus (conspicuous), and (4) less webbing in the hands (in Osteocephalus germani web reaches the antepenultimate tubercle of Finger IV, in Osteocephalus cabrerai it reaches the proximal border of the ultimate tubercle; Figs 12 and 16). Osteocephalus germani differs from other species of Osteocephalus in having a combination of a dark golden to tan golden iris, a row of indistinct tubercles in the tarsus, and areolate skin in the flanks. A golden iris with black reticulations further distinguishes Osteocephalus germani from Osteocephalus deridens , Osteocephalus oophagus , Osteocephalus planiceps , and Osteocephalus taurinus which have bronze to golden irises with black lines radiating from the pupil; iris coloration also differs in Osteocephalus carri , Osteocephalus festae , Osteocephalus heyeri , Osteocephalus subtilis , and Osteocephalus verruciger which have predominantly dark irises, and in Osteocephalus leoniae which have a bicolor iris ( Jungfer 2010; Jungfer and Lehr 2001; Lynch 2002). Osteocephalus germani differs from Osteocephalus exophthalmus , Osteocephalus fuscifacies and Osteocephalus leoniae in having abundant keratinized dorsal tubercles in males (tubercles are absent in the three last species). Skin texture in the flanks distinguishes Osteocephalus germani (areolate) from Osteocephalus mutabor (smooth). Osteocephalus inframaculatus differs from Osteocephalus germani in coloration of the ventral surfaces of hindlimbs (bold brown blotches in Osteocephalus inframaculatus are absent in Osteocephalus germani ; Jungfer 2010).

Description of holotype.

Adult male, 41.26 mm SVL, head length 12.79, head width 14.23, eye diameter 5.23, tympanum diameter 3.79, femur length 22.3, tibia length 23.1, foot length 17.97. Head narrower than body, slightly wider than long; snout rounded in dorsal view and truncate in lateral view; distance from nostril to eye longer than diameter of eye; canthus rostralis distinct and straight; loreal region concave; internarial area depressed; nostrils moderately protuberant, directed laterally; interorbital area with tiny keratinized conical tubercles, lateral margins of frontoparietals inconspicuous through skin; eye large, strongly protuberant; tympanic membrane clearly evident, large, slightly wider than high, about two thirds of eye diameter, separated from eye by ca. 85% of its diameter; tympanic annulus distinct except dorsally where it is covered by supratympanic fold; posterior end of supratympanic fold reaches arm insertion. Arm slender, axillary membrane present, reaching less than one third of arm length; four small low tubercles present along ventrolateral edge of forearm; relative length of fingers I<II<IV<III; fingers bearing large, oval discs, that of third finger about three fourths of tympanum diameter; subarticular tubercles prominent, round to ovoid except for bifid distal subarticular tubercle of Finger IV; supernumerary tubercles present; palmar tubercle small, elongated; prepollical tubercle large, flat, elliptical; prepollex enlarged; large dark keratinous nuptial excrescences covering inner surface of prepollex up to two thirds the distance between subarticular tubercle and proximal border of disk of thumb; webbing absent between fingers I and II; webbing formula of fingers II2 –-3III2½ -3-IV. Small tubercles on tibiotarsal articulation; dorsal surface of tarsus covered by tiny keratinized conical tubercles, more abundant on outer edge; minute 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 low, ovoid; subarticular tubercles single, round, protuberant; supernumerary tubercles restricted to the soles; webbing formula of toes I1-2II1-2III1+-2IV2 –- 1V. Skin on dorsum, head, and dorsal surfaces of limbs shagreen covered by conical tubercles with keratinized tips, tiny on head and limbs; skin on flanks areolate; skin on venter coarsely granular; skin on ventral surfaces of head and thighs granular, those of shanks smooth. Cloacal opening directed posteriorly at upper level of thighs; short simple cloacal sheath covering cloacal opening; round tubercles below vent; two conspicuous white tubercles ventrolateral to vent at lower level of thighs. Tongue cordiform, widely attached to floor of mouth; dentigerous processes of the vomers angular, adjacent medially, posteromedial to choanae, bearing 5 and 6 (left/right) vomerine teeth; choanae trapezoidal, oblique; vocal slits moderately long, extending diagonally from lateral end of tongue toward to the angle of snout; vocal sac indistinct above the arm and below the ear.

Color of holotype in preservative. Dorsum light brown with dark brown peripheral marks; dark brown transversal bar between orbits with fine pale borders; dorsal surfaces of forearms grayish brown with dark brown marks, dorsal surfaces of thighs, shanks, and feet grayish brown with diffuse brown transversal bars. Venter light cream with dark brown flecks on the throat and thoracic region and absent on posterior half of the body; ventral surfaces of hindlimbs and forelimbs dirty cream with dark brown flecks on the lateral borders of shanks; outer half of ventral surfaces of forearms dirty cream; sides of head brown with white subocular band extending, below tympanum, two little brown blotches below the eye; tympanic membrane dark brown and area in the periphery of tympanum light brown dorsally and grayish brown behind the tympanum; flanks grayish white, areolate region with dark brown reticulation and flecks. Iris silver with dark brown mid-horizontal line and thin black reticulations.

Color in life.

Dorsum brown with irregular dark brown marks; flanks brownish cream with dark brown spots and flecks; dorsal surfaces of thighs, shanks, and forelimbs brown with transversal dark brown bands. Venter whitish cream with brown flecks in throat; ventral surfaces of thighs tan. Iris bronze with thin black reticulations (G. Chávez field notes April 2010).

Etymology.

The new species is dedicated to our colleague German Chávez (CORBIDI), one of the best friends of PJV, for his contributions to Peruvian herpetology and collecting the type series and tissues of this new species.

Variation.

Variation in dorsal and ventral coloration of preserved specimens is shown in Figure 14. Dorsal background coloration varies from light brown to light gray; irregular dark brown marks are always present. In females, the dorsum lacks tubercles while in males tubercles are present. The single male paratype (CORBIDI 06660) differs from the holotype in having non–keratinized tubercles.

Ventral surfaces of preserved specimens (Fig. 14) are whitish cream. All the specimens have scattered dark brown flecks on the anterior half of the venter. Ventrally, limbs vary from whitish cream to tan; scant white tubercles can be present in the external edge of the forearm of males (e.g., CORBIDI 06660). The vent region is light brown or dark. Flanks are whitish cream to light gray, areolate in the anterior half and nearly smooth posteriorly. The areolate portion is completely covered by dark brown reticulation and flecks.

Snout is truncate in lateral view except for a female with rounded snout (CORBIDI 06633). Lateral head coloration varies from dull brown (CORBIDI 06633) to cream with dark brown blotches (CORBIDI 05505). The tympanic annulus is concealed dorsally and has lighter color than the background. The distal subarticular tubercle on Finger IV is bifid in all the specimens.

Adult morphometric data are summarized in Table 3.In the examined series, the largest male has a SVL of 41.45 mm and the largest female 50.76 mm; mean male SVL = 41.35 mm (n = 2, SD = 0.13), mean female SVL = 49.96 mm (n = 2, SD = 1.13).

Color in life.

Based on digital photograph of adult male CORBIDI 06660: dorsum brown with irregular dark brown marks and some scattered light green blotches; canthal region greenish brown with greenish cream subocular mark and dark labial bars; tympanum light brown; flanks light green with dark brown reticulation and dark brown blotches posteriorly; dorsal surfaces of thighs, shanks, and forelimbs brown with transversal dark brown bands and scattered light green blotches. Iris bronze with brown horizontal midline and thin black reticulations.

Based on digital photograph of adult female CORBIDI 06633: dorsum brown with few scattered irregular dark brown marks; canthal region dark brown with greenish cream subocular mark speckled by three small dark brown blotches; tympanum light brown; flanks light brown with dark brown blotches; ventrolateral region cream with fine dark reticulation; dorsal surfaces of thighs, shanks, and forelimbs brown with transversal dark brown bands. Anterior half of venter whitish cream with fine brown reticulation in throat and chest; posterior half of venter and ventral surfaces of thighs tan; iris bronze with diffuse brown mid-horizontal line and thin black reticulations.

Based on digital photograph of adult female CORBIDI 05505 (Fig. 15): dorsum green with irregular dark brown marks; canthal region green with brown mottling and white subocular mark extending to the lips as a white labial stripe along posterior half of the jaw; tympanum light brown; flanks white with dark brown reticulation and small dark brown blotches posteriorly; dorsal surfaces of thighs, shanks, and forelimbs green with transversal dark brown bands and flecks. Venter white with scattered brown flecks on throat and chest. Iris reddish gold with diffuse brown mid-horizontal line and thin black reticulations. Based on digital photograph of adult female CORBIDI 08284 (Fig. 15): dorsum light brown with irregular dark brown marks; canthal region brown and greenish white subocular mark; tympanum light brown; flanks light brown with small dark brown blotches; dorsal surfaces of thighs, shanks, and forelimbs light brown with transversal dark brown bands. Venter dull cream. Iris bronze with diffuse brown mid-horizontal line and thin black reticulations.

Distribution and ecology.

Osteocephalus germani is known from three localities in southern Peru (Fig. 6). Pongo de Manique and Comunidad Nativa de Poyentimari are in premontane forest on the Upper Urubamba River basin (vegetation types according to ONERN 1976) in the Amazonian foothills of the southern Peruvian Andes, at elevations of 670-725 m; Comunidad Nativa de Chokoriari is Terra Firme Amazonian lowland forests on the lower Urubamba River basin in the southern Peruvian Amazon lowlands, at elevation of 434 m. In Pongo de Mainique the new species was found close to rocky streams in low-hill primary forest with arboreal ferns and abundant epiphytes. At this locality, Osteocephalus germani was sympatric with Osteocephalus castaneicola and Osteocephalus mimeticus . In Comunidad Nativa de Poyentimari, Osteocephalus germani was found close to rocky streams in a step area of very wet high-hill primary forest with abundant ferns (including arboreal), epiphytes, lichens and mosses. At this locality the new species was sympatric with Osteocephalus mimeticus . In Comunidad Nativa de Chokoriari, Osteocephalus germani was found close to a black-water slow-running creek in a patch of secondary forest, surrounded by pastures for cattle and plantations. The forest was dominated by bamboo and Cecropia spp. and the creek had sandy soils covered by leaf litter. No other species of Osteocephalus were found in this locality.

All specimens were collected next to temporary pools, perching over broad leaves or on tree branches 100 to 200 cm above the ground. Many streams surround the collection sites.

Remarks.

In the phylogeny (Fig. 1), two specimens from gen bank (EF376030 from French Guiana and AY843705 from Río Jurua, Brazil) are grouped with Osteocephalus germani in a strongly supported clade (PP = 0.96) and are likely conspecific or represent one or two closely related species. The specimen from French Guiana was reported as " Osteocephalus oophagus " by Salducci et al. (2002, 2005); the specimen from Brazil was reported as " Osteocephalus cabrerai " by Faivovich et al. (2005). Both individuals appear to be misidentified.