Phrynopus capitalis, Rodriguez, Lily O. & Catenazzi, Alessandro, 2017

Rodriguez, Lily O. & Catenazzi, Alessandro, 2017, Four new species of terrestrial-breeding frogs of the genus Phrynopus (Anura: Terrarana: Craugastoridae) from Río Abiseo National Park, Peru, Zootaxa 4273 (3), pp. 381-406 : 389-392

publication ID	https://doi.org/ 10.11646/zootaxa.4273.3.4
publication LSID	lsid:zoobank.org:pub:ADC167F6-C3A0-43EC-8636-F65F5ABE85AF
DOI	https://doi.org/10.5281/zenodo.6033268
persistent identifier	https://treatment.plazi.org/id/ED0DF709-FFB6-7543-9EB4-73EB95EFF932
treatment provided by	Plazi
scientific name	Phrynopus capitalis
status	sp. nov.

Treatment

Phrynopus capitalis new species

( Figures 6 View FIGURE 6 B, 7, 8, 9A)

Holotype. AMNH 134158 About AMNH , an adult male ( Figures 6 View FIGURE 6 B, 7) obtained at Lake Manachaqui (7°41’47” S, 77°30’56” W, 3600 m), ca. 14 km airline NE Pataz, Provincia Pataz, Departamento (Región) La Libertad, Peru ( Figure 1 View FIGURE 1 ), collected on 3 July 1989 by L.O. Rodríguez. GoogleMaps

Paratopotypes. MUSM 8959, a gravid female ( Figure 8 View FIGURE 8 ), and MUSM 8957, a juvenile, collected with the holotype.

Diagnosis. A moderately-sized Phrynopus with the following characteristics: (1) skin on dorsum and flanks fleshy and tuberculated, densely covered with minute round tubercles, bearing low warts especially above the insertion of the forelimbs and dorsolaterally; skin on throat, chest and belly coarsely areolate, ventral surface of thighs coarsely areolate; discoidal fold absent, thoracic fold evident; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus absent; supratympanic fold swollen (similar to a parotoid gland); (3) snout short, nearly rounded in dorsal view, sloping and rounded in profile; nasals large, frontoparietals in contact; (4) upper eyelid without tubercles; width of upper eyelid narrower than IOD; cranial crests absent; (5) dentigerous processes of vomers absent; (6) males with gray nuptial pad on thumb, lacking vocal slits and vocal sac; (7) Finger I slightly shorter than Finger II; tips of digits bulbous, terminal phalanges T-shaped ( Figure 9 View FIGURE 9 A); (8) fingers bearing basal webbing and fleshy lateral fringes; (9) ulnar and tarsal tubercles absent; (10) heels lacking tubercles; inner tarsal fold absent; (11) inner metatarsal tubercle rounded, about 1.5 times larger than rounded outer metatarsal tubercle; supernumerary plantar tubercles absent (12) toes bearing lateral fringes; basal webbing present; Toe V longer than Toe III; toe tips rounded, bulbous, about as large as those on fingers; terminal phalanges with lateral processes (as in Figure 9 View FIGURE 9 A) (13) in life, dorsal surfaces are brownish tan with small, dark brown reticulate spots having the appearance of a labyrinth; areolation on ventral surfaces are grayish cream in a darker ground color; in preservative, flanks are blueish black or black; (14) SVL in single female 35.6 mm, in single male 23.2 mm.

The new species cannot be confused with any other described species of Phrynopus because of its very robust and prominent head (HW/SVL = 40 %), swollen tympanic area, absence of tympanic annulus, thick lateral fringes on all digits, and its distinctive dorsal coloration pattern with small brown reticulated spots. Phrynopus capitalis differs from P. auriculatus , P. montium and P. peruanus by lacking a visible tympanic annulus. Phrynopus anancites , P. auriculatus , P. bracki , P. dagmarae , P. interstinctus , P. kauneorum , P. kotosh , P. lechriorhynchus , P. nicoleae , P. peruanus , P. vestigiatus all differ (character condition for P. capitalis in parentheses) by having the dentigerous processes of vomers (absent). Phrynopus badius , P. curator , P. horstpauli , and P. thompsoni have discontinuous or ridge-like dorsolateral folds (absent). Of the species lacking a tympanum, the dentigerous processes of vomers and dorsolateral folds or ridges, P. barthlenae has similar size, coarsely tuberculate dorsal skin, and lateral fringes on digits, but differs from P. capitalis by lacking warts (present), by bearing a dark canthal and postorbital stripe (absent), by having small nasals (large) and widely separated frontoparietals (large nasals, frontoparietals in contact), and distinctive black dorsal blotches (labyrinth-like short spots on dorsal surfaces). Phrynopus bufoides is most similar to P. capitalis , but differs by having a discoidal fold (absent), a broad supratympanic fold (swollen), by lacking lateral fringes on toes (present), and by bearing prominent dorsal isolated and elongated warts forming discontinuous ridges (coarsely areolate dorsal skin with warts and minute tubercles), and black mottling on venter (uniform ventral coloration). Phrynopus chaparroi lacks lateral fringes on digits (present), has a much darker coloration without dorsal pattern (tan with brown pattern); P. daemon is readily distinguished by having subacuminate snout in lateral view (rounded), short supratympanic fold (swollen), no dorsal pattern (short labyrinth-like lines) and males having prominent subgular vocal sac (absent), and by lacking nuptial pads (present). Phrynopus heimorum has a slightly tuberculate dorsum (fleshy and tuberculate) and light reticulations especially on flanks and venter (dorsum with labyrinth-like small spots, venter uniform), and lacks lateral fringes on all digits (present); P. juninensis has smooth dorsal and ventral skin (coarsely areolate). Phrynopus miroslawae differs by lacking lateral fringes on digits and basal webbing on toes (present), by bearing dorsolateral, occipital and supratympanic folds (absent, supratympanic fold swollen), by having a row of subconical tubercles on outer edge of tarsus (absent), and by being smaller with the single known female having SVL = 29.2 mm (SVL = 35.6 mm). Phrynopus pesantesi lacks lateral fringes on digits (present), has a blunt snout (sloping to rounded in lateral profile) and smaller head with HW/SVL = 36% (40% in P. capitalis ), a tuberculate dorsum (tuberculated bearing low warts), and olive and gray mottled and yellow blotches on groin, ventral surfaces of throat, upper arms, forearms, thighs, shanks and venter. Phrynopus valquii differs from P. capitalis by lacking lateral fringes or basal webbing in toes and fingers, by bearing canthal and supratympanic stripes (absent), and by having dorsal skin shagreen with scattered tubercles (coarsely areolate). Phrynopus tautzorum has a smooth dorsum with scattered tubercles (fleshy and tuberculate), lacks lateral fringes on digits (present), and has Toe V slightly shorter than Toe III (Toe V longer than Toe III). The following species of the Pristimantis orestes group with reduced finger tips may externally resemble Phrynopus capitalis : Pristimantis atrabracus , P. chimu , P. cordovae , P mariaelenae , P. melanogaster , P. pataikos , P. pinguis , P. seorsus , P. simonsii , P. stictoboubonus , P. stipa . All of these except P. simonsii and P. attenbouroughi , have a visible tympanum ( P. atrabracus , P. chimu , P. cordovae , P. mariaelenae , P. pinguis ), or the tympanic annulus is weakly defined or concealed beneath the skin ( P. melanogaster , P. pataikos , P. stictoboubonus , P. seorsus , P. stipa ). The other two members of the group occurring in Peru, P. corrugatus and P. ventriguttatus have well developed pads on all digits and cannot be confused with Phrynopus capitalis . Pristimantis simonsii lacks a tympanum, dentigerous processes of vomers, and bears lateral fringes on digits, as does Phrynopus capitalis , but differs by having relatively larger nasal bones, by lacking the swollen supratympanic fold, by having smoother dorsal skin (coarsely tuberculate with warts) and by bearing some dorsal dark blotches (absent). Pristimantis attenbouroughi differs by bearing the dentigerous processes of vomers (absent), by having dorsal skin shagreened with low scattered tubercles (coarsely areolate with large rounded warts), a dark canthal and supratympanic stripe (absent, supratympanic fold swollen), and by being smaller with males SVL = 14.6–19.2 mm (single male SVL = 23.2 mm; Lehr and von May 2017).

Description of holotype. Head distinct, narrower than body, wider than long, (HW 133% of HL; HW 40% of SVL; snout nearly rounded in dorsal view, sloping to rounded tip in lateral profile; snout moderately short; E-N 70% of eye diameter; nostrils small, weakly protuberant, dorsolateraly directed; canthus rostralis slightly curved in dorsal view, rounded in profile; loreal region slightly concave, lips weakly flared; interorbital area slightly convex, no cranial crests; upper eyelid without enlarged tubercles; EW 90% of IOD in male specimen. Tympanic membrane and annulus absent; tympanic region with several irregular fused tubercles or warts, some of which are arranged in a swollen fold (similar to a parotoid gland), extending from the posterior margin of the eye to the shoulders. Oval to elongate, large tongue, weakly notched; choanae small, round, well separated and anterolaterally situated; dentigerous processes of vomers absent. Vocal sac and vocal slits absent.

Skin of dorsum irregularly granular bearing some warts and small ridges, especially in the scapular area and on the lower back, bearing scattered very small tubercles; skin of venter and throat coarsely areolate; two large postrictal tubercles under the swollen supratympanic fold; a low elongate tubercle on the interorbital area; metacarpal tubercles large, oval and low; outer areas of forelimbs bearing some elongate warts; inner palmar tubercle nearly rhomboidal, thenar round, small slightly shorter than palmar; fingers bearing thick lateral fringes, finger tips rounded and slightly bulbous; first finger slightly shorter than second; digital pads present; subarticular tubercles round, conical in profile ( Figures 6 View FIGURE 6 B, 9A); THL 38% of SVL; two metatarsal tubercles, round and large, inner one and a half the size of outer; some supernumerary plantar tubercles ( Figure 6 View FIGURE 6 B). Toes bearing lateral fringes and basal webbing; toe tips bulbous, emarginated; Toe V longer than Toe III.

Color in life. Dorsal surfaces were brownish-gray on the scapular region on the holotype and brownish-tan bluish black on the flanks; the venter was yellowish gray. Iris cocoa brown. Nuptial pads black.

Color in preservative. Dorsal surfaces are brownish-tan with small dark brown spots appearing like a labyrinth; flanks are black with some brown on the warts ( Figure 7 View FIGURE 7 ). Groin with black reticulations; posterior thigh black or brownish-tan, with the same dorsal pattern. Ventral surfaces grayish cream on the areoles with darker grayish ground color.

Measurements in mm (data for male holotype followed by female paratopotype in parentheses). SVL 23.2 (35.9), HL 7.0 (13.6), HW 9.3 (14.3), TL (9.0 (12.0), THL 9.1 (13.0), FL 9.4 (13.9), IOD 2.9 (3.6), EW 2.6 (3.3), E-N 2.1 (2.8), IND 1.6 (2.7), ED 3.0 (3.7).

Variation. Female MUSM 8959 is larger in size (35.6 mm), and EW is narrower than the IOD (wider in the holotype). The two postrictal tubercles are also present. Coloration is the same as in the holotype. In life ( Figure 8 View FIGURE 8 ), bluish black coloration under arms and above groin, more conspicuous than in male holotype.

Etymology. The specific name capitalis is a Latin adjective meaning “pre-eminent” and is used in reference to the distinctive large head of the species

Distribution and ecology. Phrynopus capitalis is known only from the type locality in the Manachaqui Valley, where it co-occurs with Phrynopus valquii and an undescribed species of Pristimantis . This valley is covered with wet montane grasslands and contains a glacial lake that feeds a tributary of the Marañon river ( Figures 2 View FIGURE 2 , 3 View FIGURE 3 C). The three specimens were found within a 100 m 2 area in the wettest flat area of this valley, among dense moss vegetation. This area lays outside of the Río Abiseo NP, but P. capitalis might be present in areas of the same habitat within the park.