Phrynopus dumicola, 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 : 392-396

Phrynopus dumicola new species

( Figures 6 C, 9B, 10, 11)

Holotype. MUSM 33102 , a gravid female ( Figures 6 C, 10) obtained at Pampa del Cuy (7°39'49"S, 77°28'43"W; 3470–3550 m), ca. 18 km airline NE Pataz, Parque Nacional del Río Abiseo, Distrito Pataz, Provincia Mariscal Cáceres, Departamento (Región) San Martín, Peru, collected on 19 July 2000, by A. Catenazzi and R. von May.

Paratopotypes. Eleven adult males, one adult female and one juvenile female: males MUSM 33092 and 33118, collected on 18 and 19 July 2000 by A. Catenazzi and R. von May; female MUSM 3819, collected on 1 July 1987; males MUSM 3815, 3817, AMNH 134167 (cleared and stained), and juvenile female MUSM 3820, collected on 19 July 1987; males AMNH 134149–50 and 134151 (cleared & stained), collected on 29 July 1988; males MUSM 9078–79, collected on 3 August 1990, all collected by L. O. Rodríguez.

Paratypes. Two adult males, one adult female and one juvenile male: adult male AMNH 134152 collected in elfin forest between Pampa del Cuy and Puerta del Monte (ca. 7°40'S,77° 27'W; 3350 m) on 19 July 1988; adult female MUSM 3818, adult male KU 220917, and juvenile male USNM 299788, collected at Puerta del Monte, ca. 20 Km airline NE Pataz, (7°39'29.64"S, 77°28'12.06"W; 3225 m) on 18 July 1987 by L. O. Rodríguez.

Diagnosis. This species of Phrynopus has: (1) skin on dorsum shagreen or rugose with low scattered tubercles, skin on flanks areolate, skin on throat smooth, that of chest and venter areolate but not as coarsely as on flanks, ventral surface of thighs coarsely areolate; discoidal fold absent, thoracic fold present; dorsolateral ridges present; (2) tympanic membrane and tympanic annulus absent, moderate supratympanic fold present; (3) snout short, rounded in dorsal and lateral views; (4) upper eyelid without conical or rounded tubercles; EW narrower than IOD; cranial crests absent; (5) dentigerous processes of vomers absent; (6) males lacking vocal slits and nuptial pads; (7) Finger I slightly shorter or equal in length than Finger II; tips of digits rounded, terminal phalanges knob-shaped; (8) fingers without lateral fringes; (9) ulnar and tarsal tubercles absent; (10) heels lacking tubercles; inner tarsal fold absent; (11) inner metatarsal tubercle ovoid, about 1.5 times larger than rounded outer metatarsal tubercle; supernumerary plantar tubercles absent; (12) toes without lateral fringes; basal webbing absent; Toe III longer than Toe V; toe tips rounded, about as large as those on fingers; (13) in life, dorsum dark brown, reddish brown or olive green with irregular darker blotches, dark brown post orbital stripe present; throat varying from pearly white to pale salmon or creamy yellow, chest and belly dull brown to creamy white or bluish white with or without brown irregular blotches; ventral surfaces of forearms and thighs brown or salmon, ventral surface of hands and feet dark brown with irregular creamy white flecks or blotches; groin dark brown, or reddish brown with pearly white flecks; (14) SVL in females 22.3–25.3 mm (n = 5), in males 18.3–22.55 mm (n = 6).

Seven other species of Phrynopus have aerolate skin ventrally, lack the tympanic annulus and dentigerous processes of vomers, but can be distinguished from P. dumicola by a combination of the following characters (character condition for P. dumicola in parentheses). Phrynopus barthlenae bears lateral fringes on all digits (absent), has basal webbing on toes and nuptial pads in males (absent), and a dorsal pattern (absent). Phrynopus bufoides is larger with females up to 33.6 mm SVL (25.2 mm) and wears large warts on dorsum (absent), whereas P. heimorum has a distinctive red coloration on ventral side of limbs (ventral coloration not red) and T-shaped terminal phalanges (not expanded). Phrynopus daemon has a weakly areolate skin (areolate), wears lateral fringes on toes (absent), and males have a subgular vocal sac (absent). Phrynopus horstpauli is larger with females up to 39.7 mm SVL, possesses a longer snout, toe V>III (toe III>V), and Finger I is much shorter than II. Phrynopus oblivius has a longer snout (E-N/ED 68-69% in P. oblivious , 60-61% in P. dumicola ) smooth dorsal skin, bears ulnar tubercles (absent), has dark labial stripes (absent) and light small spots in venter (absent). Phrynopus valquii also lacks a tympanic annulus and the dentigerous processes of vomers, but differs by having a shorter and wider head, by lacking dorsolateral folds (discontinuous), by having shorter digits and bulbous digital tips (round tips), by bearing irregularly T-shaped terminal phalanges (knob-shaped; see Fig. 9 B and 9D), and by having lighter dorsal coloration, and frontoparietals completely in contact (moderately separated).

Phrynopus chaparroi , P. paucari and P. thompsoni also share most characters with P. dumicola , but P. chaparroi is larger, with females up to 32.2 mm in SVL, males wear nuptial pads (absent), bulbous toe tips (round), and a dark venter with irregular, diffuse white to gray blotches (ventral coloration light and lacking a pattern). Phrynopus paucari possesses a head that is longer than wide (HL <HW), has a discoidal fold (absent) and is bright greenish-yellow on the hands and the inner side of limbs and flanks, with fine reticulations on venter in live specimens (dull coloration ventrally, sometimes with salmon or dull pink on inner side of limbs). Phrynopus thompsoni has a longer snout, wears longitudinal dorsal rows of low tubercles with pustules (dorsum shagreen with low tubercles), lateral fringes on toes (absent), has Toe III = V (III> V) and many distinct brown spots on creamy tan venter (venter without any spots).

Other species superficially resemble Phrynopus dumicola , but differ by having the dentigerous processes of vomers. Of these, P. kotosh further differs by bearing small tubercles on heels (absent) and by having transversal bars on thighs and lower frank (absent), whereas P. bracki differs by having a concealed tympanum (absent) and a very short first toe (longer than wide).

Species of the Pristimantis orestes group with reduced digital pads may externally resemble Phrynopus dumicola ; those are P. atrabracus , P. chimu , P. cordovae , P mariaelenae , P. melanogaster , P. pataikos , P. pinguis , P. seorsus , P. simonsii , P. stictoboubonus , and P. stipa . All of those species, except P. simonsii , 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 orestes group occurring in Peru, P. corrugatus and P. ventriguttatus bear well developed pads on all digits and cannot be confused with Phrynopus dumicola . Pristimantis simonsii lacks a tympanum and dentigerous processes of vomers, as in Phrynopus dumicola , but differs by bearing lateral fringes and basal webbing (absent), by having relatively larger nasal bones, a shorter head and presenting black blotches on groins (no marks). Pristimantis attenbouroughi is slightly smaller than dumicola , males SVL to 19.2 mm, females SVL to 23.0 (males SVL to 22.6mm, females SVL to 25.3 mm), bears the dentigerous processes of vomers (absent), has a relatively longer snout with EN 70% of ED (EN 60% of ED), heel with a small conical ulnar and tarsal tubercles (absent), a short cloacal sheat (absent) and lighter dorsal coloration with stripes on low flanks (absent). Pristimantis stipa externally resembles Phrynopus dumicola , but differs by bearing a supratympanic fold and tympanic annulus (absent in P. dumicola ), by bearing the dentigerous processes of vomers, bulbous digital tips (round), prominent subarticular tubercles (weakly defined), and by having white mottling on venter, and white nuptial pads in dorsal and medial surfaces of thumbs in males (absent).

Description of holotype. Head narrower than body, wider than long; HW 103% of HL; HW 37% of SVL; snout nearly rounded in dorsal view, rounded in lateral profile; canthus rostralis rounded to moderately sharp in lateral view; nostrils small, slightly protuberant, laterally directed; loreal region concave, sloping; snout short, E-N 86% of eye length; interorbital region flat, no cranial crests; EW 83% of IOD; tympanic membrane and annulus absent; supratympanic fold short, not prominent bordered by a dark stripe; tongue oval, not notched and one-third free posteriorly; choanae small, round, not hidden by palatal shelf of maxillae; dentigerous processes of vomers absent; skin of dorsum shagreen with low scattered tubercles, low postocular folds, discontinuous dorsolateral folds; some low small supraorbital tubercles; skin of venter coarsely areolate extending to the flanks and postrictal area; skin of throat areolate; moderate thoracic fold present, no discoidal fold; no ulnar tubercles; weakly defined palmar tubercle, palmar slightly smaller than thenar tubercle; fingers lacking lateral fringes; Finger I equal to Finger II.

Hind limbs slender, short, TL 35% of SVL; upper surfaces of hind limbs smooth; posterior and ventral surfaces of thighs coarsely areolate; heel and outer surface of tarsus lacking tubercles; two metatarsal tubercles, inner oval; outer rounded, half size of inner; no supernumerary plantar tubercles; slightly areolate plantar surface; toes lacking lateral fringes and basal webbing; toe tips rounded, lacking marginal grooves, about as large as those on fingers; Toe III longer than Toe V ( Fig. 4 C).

Color in life. Overall dark brown dorsally and ventrally, with some light reddish areas on throat, chest and upper venter. Iris bronze, turning darker towards the rounded pupil.

Color in preservative. Dark brown to brown dorsally and laterally, with a visible supratympanic stripe. Venter brown to light brown with some darker spots.

Measurements of the holotype (in mm). SVL 23.3, HL 8.3, HW 8.6, TL 8.5, THL 9.2, FL 10.8, IOD 2.7, EW 2.3, IND 2.5, ED 3.2, E-N 2.7.

Variation. Coloration pattern in some specimens (MUSM 3819 and 33116) includes a middorsal line that extends onto back of thigh. Some specimens (e.g., MUSM 3815) have lighter areas on throat and venter. In darker specimens, the supratympanic fold bordered by a dark stripe is barely visible (MUSM 9 0 78 and 9079). The skin of dorsum is usually shagreened, forming discontinuous dorsolateral ridges and occipital low folds.

Etymology. The specific name dumicola is a Latin adjective meaning "a dweller in thickets" in reference to the habitat of forest patches bordering the tree line, inhabited by the species.

Distribution and ecology. This species is only known from Pampa del Cuy, Río Abiseo NP. Most specimens were found in mosses on the ground of patches of elfin forest patches ranging from 0.01 to 5 ha in size. These elfin forests grow on rocky substrates surrounded by wet montane grassland along a linear distance shorter than 3 km in the Pampa del Cuy Valley ( Figure 3 B), from 3350 to 3550 m. The holotype was found in the grassland adjacent to forest patches, during a rainy night. Phrynopus dumicola was the only species of frogs encountered in leaf-litter plots in these elfin forest, while the surrounding grasslands contained other species of Pristimantis and Gastrotheca . We found at least one P. dumicola specimen in every forest patch. We found some eggs (4.5 mm in diameter) with adult specimens on 19 July 1987, suggesting parental clutch attendance. Most P. dumicola had trombiculid mites under the skin, especially in the ventral area and limbs.

Figures

FIGURE 3. Habitats of species of Phrynopus in the Río Abiseo NP and surrounding area. (A, B) habitat of P. capitalis sp. nov. and P. valquii in the Manachaqui valley, 3500 m, Department of La Libertad, near the border of Río Abiseo NP. (C) Pampa del Cuy valley, showing forest patches on the slopes; habitat of P. dumicola sp. nov.. (D) View over the Montecristo Valley, from El Mirador, habitat of P. personatus sp. nov. Photographs by A. Catenazzi.

FIGURE 4. Holotype of Phrynopus anancites sp. nov., male MUSM 33168 (SVL = 25.3 mm), showing (A) head in lateral view and (B, C) body in dorsal and ventral views. Specimen collected at Ventanas, 3820 m, near the entrance of Río Abiseo National Park, San Martín, Peru. Photographs by Morris Flecks.

FIGURE 6. Palmar and plantar surfaces of the new species of Phrynopus: (A) P. anancites sp. nov. (MUSM 33168), (B) P. capitalis sp. nov. (AMNH 134158), (C) P. dumicola sp. nov. (MUSM 33102), and (D) P. personatus sp. nov. (MUSM 33096). Photographs by M. Flecks (A, C and D) and A. Catenazzi (B).

FIGURE 9. Condition of terminal phalanges of Finger III, left hand: T-shaped in (A) Phrynopus capitalis sp. nov. (MUSM 8959), and knob-shaped in (B) P. dumicola sp. nov. (USNM 299788), (C) P. personatus sp. nov. (MUSM 3813), (D) P. valquii (MUSM 3823). Drawings by F. Chang.