Tepuihyla shushupe, Ron, Santiago R., Venegas, Pablo J., Ortega-Andrade, H. Mauricio, Gagliardi-Urrutia, Giussepe & Salerno, Patricia E., 2016

Ron, Santiago R., Venegas, Pablo J., Ortega-Andrade, H. Mauricio, Gagliardi-Urrutia, Giussepe & Salerno, Patricia E., 2016, Systematics of Ecnomiohylatuberculosa with the description of a new species and comments on the taxonomy of Trachycephalustyphonius (Anura, Hylidae), ZooKeys 630, pp. 115-154 : 129-136

Taxon classification Animalia Anura Hylidae

Tepuihyla shushupe sp. n.

Ecnomiohyla tuberculosa : Venegas and Gagliardi-Urrutia (2013). Species misidentification in their Figures 8B and 8G.

Holotype.

(Figs 3-4, 6, 10C) CORBIDI 12513, adult male from Peru, Loreto department, Maynas province, head waters of rivers Ere and Campuya, Putumayo River basin (1.6790°S, 73.7197°W), 145 m above sea level, collected by P. J. Venegas and G. Gagliardi-Urrutia on 19 October 2012.

Diagnosis.

In this section coloration pertains to preserved specimens unless otherwise noted. A large-sized Tepuihyla differing from other species in the genus by the following combination of characters: (1) SVL in males 85.4 mm (n = 1), females unknown; (2) skin on dorsum coarsely tuberculate, covered by small tubercles intermixed with large tubercles; tubercles without keratinized tips; (3) skin on flanks similar to skin on dorsum; (4) webbing between fingers extensive but without reaching the proximal border of disks; hand webbing formula I2-2II1- 2III 1½ -1+IV (Fig. 7C); webbing between toes extensive, reaching the proximal border of the disks on at least three toes; foot webbing formula I1-1+II0+-1+III0+- 1½IV1½ - 0+V (Fig. 7D); (5) dorsal coloration in life brownish green, anterior and posterior surfaces of thighs and hidden surfaces of shanks yellowish orange; (6) ventral coloration whitish gray and webbing between fingers and toes pale orange; (7) suborbital mark absent, clear labial stripe present, faint and with some brown spots; (8) coloration on flanks similar to dorsal coloration except for greenish yellow axilla and groins and whitish gray ventrolateral area; (9) skin on upper surface of head not co-ossified with underlying cranial element, cranial crest slightly exostosed; (10) in life, bones green; (11) small triangular dermal flaps form serrate fringe along the ventrolateral margin of the forearm and along the outer edge of Finger IV; heels bearing two enlarged fleshy conical tubercles surrounded by few smaller round tubercles; small triangular dermal flaps form serrate fringe along ventrolateral margin of tarsus and outer margin of Toe V; (12) in life, iris reddish peripherally turning cream towards to the center, with irregular black reticulations; (13) vocal sac is white, single and subgular, (14) juveniles unknown; (15) larvae unknown.

Tepuihyla shushupe differs from all species of the genus (in parenthesis), except Tepuihyla tuberculosa , in being larger, having extensive hand webbing (basal webbing) and a serrate fringe along the ventrolateral margin of the forearm (absent). Other differences are listed on Table 4.

Tepuihyla shushupe is most similar to Tepuihyla tuberculosa . It differs from Tepuihyla tuberculosa (character states in parenthesis) in having, in life, a cream iris with red periphery (iris cream without red periphery), dorsum covered by small tubercles intermixed with abundant large tubercles (dorsum covered by small tubercles intermixed with few large tubercles); in preservative, dorsum cream with a brownish mantle (dorsum light cream or cream suffused with a coppery hue) and the posterior border of large dorsal tubercles dark brown (tubercles brownish cream or creamy coppery). In coloration, size, and texture of dorsal skin Tepuihyla shushupe resembles the Amazonian Trachycephalus resinifictrix and Trachycephalus cunauaru . It can be distinguished from both species by the presence of serrate fringes on the limbs and tubercles in the lower jaw (both absent in Trachycephalus ) and the iris having red periphery and lacking a vertical black bar below the pupil (iris without red and with a vertical back bar below the pupil in both Trachycephalus ).

Description of the holotype.

Adult male (CORBIDI 12513), 85.4 mm SVL, head length 26.6, head width 28.9, eye diameter 7.7, tympanum diameter 5.5, femur length 35.9, tibia length 46.9, foot length 35.9. Head rounded in dorsal view, wider than long (HL/HW = 92%); snout truncate in dorsal view and slightly protruding in profile; nostrils are directed laterally, near tip of the snout; top of head tuberculate; canthus rostralis concave in dorsal view, straight in profile; skin on upper surface of head not co-ossified with underlying cranial elements, cranial crests slightly exostosed; loreal region concave; skin on dorsal surface of head, body and limbs coarsely tuberculate, tubercles are either small or large and lack keratinous spines; lower eyelid pigmented; distinct supratympanic fold formed by a row of round tubercles running from midpoint of posterior margin of eye; tympanum prominent, opaque, smooth, 71% of eye diameter, separated from eye by 3.8 mm; small triangular dermal flaps form serrate fringe along the ventrolateral margin of the forearm and the outer edge of Finger IV, fringe more conspicuous on forearm that on finger; hands moderate in length (HAL/SVL = 31.2%); Finger lengths I <II <IV <III, terminal disk on Finger I 63.2% of diameter of disks on Fingers II–IV; diameter of Fingers II-IV ~1.2 times tympanum diameter; distal subarticular tubercles on Fingers I–III large and rounded, on Finger IV larger and bifid; supernumerary tubercles indistinct; prepollical tubercle large, obtuse and elliptical; bony prepollical projection absent; keratinous nuptial pad with dark epidermal projections covering the surface of the thumb up to the distal edge of its subarticular tubercle; fingers considerably webbed although web does not reach base of disks; webbing formula I2-2II1- 2III 1½-1¼ IV; legs relatively long and slender (tibia length 54.8% of SVL), thigh length 35.9 mm; heels bearing two enlarged conical tubercles surrounded by few lower round tubercles; fleshy, dermal flaps form serrate fringe along ventrolateral margin of tarsus and outer margin of Toe V extending to base of disk, scallops are deeply incised, largest on tarsus, smaller along toe; outer metatarsal tubercle small, inner metatarsal tubercle moderately large, ovoid, and flat; toe lengths I <II <III = V <IV; disks on toes ~70% of diameter of disks on fingers, equal between Toes III–V; subarticular tubercles round; supernumerary tubercles present, ill defined; webbing in toes extensive, reaching the base of disks on at least three toes; webbing formula I1-1II1- 1¼III¾ -1IV1- ¾ V; gular skin smooth with small pointed tubercles along the jaw, venter finely granulate, weak granulation on undersides of arms and finely on ventral surfaces of thighs, smooth skin on anterior surfaces of thighs and ventral surfaces of legs; cloacal opening directed posteriorly at upper-level of thighs, some indistinct small tubercles on the upper edge of vent, skin under the vent covered by small flattened but conspicuous tubercles with two enlarged prominent fleshy tubercles posterolateral to the vent (one on each side); tongue cordiform; vomerine ridges transverse, oblique in the middle, narrowly separated medially, placed between the posterior margins of the large subrectangular choanae; vomerine teeth 6 to 11; vocal slits present.

Color of holotype in life

(Figs 3, 10). Based on digital photographs. Dorsal and lateral surfaces of body and limbs pale green; posterior edge of each enlarged tubercle, dark brown; lips paler than the rest of head; lower flanks, forearms, tibia, and tarsus white; dorsal surfaces of hindlimbs with narrow dark brown transversal stripes; axillar region, groins, anterior and posterior surfaces of thighs, and hidden surfaces of shanks greenish yellow; dorsally, webbing between fingers and toes pale greenish yellow with scattered dark brown marks, ventrally greenish orange. Venter whitish gray with an orange tone on belly and ventral surfaces of thighs; ventral surfaces of arms and elbows greenish yellow; ventral surfaces of tibia, tarsus, palms, fingers, soles, toes and discs water green; limb bones green; iris cream with red periphery and irregular black reticulations.

Color of holotype in preservative

(Figs 4 A–B, 5 C–D). Dorsal surfaces of head, body and forelimbs, including fingers, are tan; large dorsal tubercles have a dark brown margin; dorsal surfaces of hindlimbs and toes dirty cream bearing scattered light tubercles with dark brown margins and narrow dark brown transversal stripes; posterior surfaces of thighs light cream; dorsal surfaces of discs on fingers and toes are light cream with dark brown marks. Venter is whitish cream.

Advertisement call.

Quantitative characteristics of the advertisement call Tepuihyla shushupe (CORBIDI 12513) are detailed in Table 3. The call consists of a cackle of short notes repeated at a fast rate with amplitude modulation (Fig. 6). Note amplitude and note rate increases markedly along the first half of the call, decreasing again at the end. The call has two harmonics, the first harmonic has slightly more energy than the second (Fig. 6). Fundamental frequency of individual notes ranges from 404.3 to 585.9 Hz (mean = 500.08, SD = 28.3). The dominant frequency of the entire call is 515.6 Hz. Midway through the call, nearly 25% of the notes have the greatest energy in the second harmonic (range 1016.6 to 1210 Hz).

Distribution and natural history.

Tepuihyla shushupe is only known from the type locality in the headwaters of rivers Ere and Campuya, at an elevation of 145 m, in the Putumayo river basin near the boundary between Peru and Colombia. According with Vriesendorp (2013), the type locality consists of a complex of forest terraces, at elevations between 90 and 170 m above sea level, with a canopy reaching 35 to 40 m; terraces have heavy loads of leaf litter (~50 cm deep) and a dense mat of fine roots; the depressions between terraces have small palm swamps (~10 m wide) of Oenocarpus bataua ; the soil varies between sandy and clayey; most streams have a muddy bottom, few have gravel and sand, and one has big cobbles.

The holotype was calling at the base of a big tree inside a narrow hole, 150 cm above the ground. The hole had 30 cm of height and had water accumulated. The frog had most of its body submerged (Fig. 10C) and the top of its head was covered by flies (probably Corethrella midges). We recorded its call and immediately made playbacks. The male answered by calling quickly and perching on the hole entrance. We detected at least six individuals during 18 hours of visual encounter surveys (0.375 individuals/hour,) in areas nearby where the holotype was found (primary forest). All individuals were detected by their advertisement calls and none could be collected. No females, amplectant pairs, clutches, or tadpoles have been observed.

Etymology.

The specific epithet is a noun in apposition. The word shushupe is used by native people to refer to the bushmaster Lachesis muta ( Squamata : Viperidae ), the largest viper in the Americas. Our field assistants in Ere river, Alpahuayo Mishana (Peru) and Juyuintza (Ecuador) believed that the advertisement calls of Tepuihyla shushupe and Tepuihyla tuberculosa were produced by Lachesis muta . The belief that Lachesis muta can sing seems to be widespread among hunters, colonists, and indigenous people from the Amazon basin ( Lamar 1998). The association of the calls from Tepuihyla with Lachesis muta by people on widely separated localities in Amazonian Peru and Ecuador deserves investigation.

Conservation status.

Tepuihyla shushupe is only known from a single individual collected at the type locality. Calling behavior suggests that Tepuihyla shushupe breeds on tree holes and is a canopy dweller. The detection of the calls of six additional individuals at the type locality suggests that the species can be relatively abundant. However, the species may be difficult to observe in ground-level surveys. Given the uncertainty in its population status, we suggest that its Red List category is Data Deficient according to IUCN (2001) guidelines.

Remarks.

The juvenile specimen (USNM 193866) from Rio Querari (Amazonas, Brazil) reported as Ecnomiohyla tuberculosa by Duellman (1974) probably correspond to Tepuihyla shushupe , but a taxonomic validation is needed to confirm its identity.