Phrynopus personatus, 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 : 396-401

Phrynopus personatus new species

( Figures 6 D, 9C, 12, 13)

Holotype. MUSM 33096 , a gravid female ( Figures 6 D, 12) obtained near El Mirador (7°39'50"S, 77°26'55"W; 2890 m), ca. 20 km airline NE Pataz, Distrito Huicungo, Provincia Mariscal Cáceres, Departament (Región) of San Martín, Peru, collected on 26 July 2000 by A. Catenazzi and R. von May.

Paratopotypes. MUSM 33166, an adult male collected with the holotype.

Paratypes. Five adult males and six adult females, all collected near the type locality, but at different elevations: males MUSM 33121 (7°39'33"S, 77°28'07"W, 3260 m), MUSM 33109 (7°39'36"S, 77°27'46"W, 3180 m); females MUSM 33116 (7°39'36"S, 77°27'46"W, 3160 m), MUSM 33091 and 33099 (7°39'48"S, 77°28'26"W, 3110 m), MUSM 33100 (7°39'43"S, 77°27'36"W, 3150 m), collected on 21–22 July 2000 by A. Catenazzi and R. von May; female MUSM 3813 (7°39'45.92"S, 77°27'11.04"W, 3000 m), male AMNH 134153 and female MUSM 3811 (2930 m) collected on 23 July 1988 by L.O. Rodríguez; and males MUSM 3810 and 3812 (7°39´48” S, 77° 28´26” W, 3100 m) collected on 26 July 1988 by L. O. Rodríguez.

Diagnosis. This species of Phrynopus has: (1) skin on dorsum shagreen with low scattered round tubercles on dorsum and flanks, skin on flanks areolate; skin on throat, chest and belly finely areolate, ventral surface of thighs coarsely areolate; discoidal fold absent, fine subgular and thoracic folds present; dorsolateral folds and an X shaped medial ridge usually present; (2) tympanic membrane and tympanic annulus absent; (3) snout rounded in dorsal and lateral views; (4) upper eyelid bordered by a dermal ridge and bearing some low rounded tubercles; width of upper eyelid narrower than IOD; cranial crests absent; (5) dentigerous processes of vomers of vomers absent; (6) males lacking vocal slits and nuptial pads; (7) Finger I slightly shorter than Finger II; tips of digits rounded, terminal phalange knob-shaped ( Fig. 9 c); (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; Toe V slightly longer than Toe III; 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 canthal and post orbital stripe marking always a face mask; throat and ventral surfaces of forearms and thighs reddish brown to red-wine, ventral surface of hands and feet lighter or dark brown with scattered creamy white flecks or blotches topping tubercles; low venter and groin region dark brown, or black with sharp defined round white blotches, yellow in life, sometimes extending onto the shank and also present behind the thigh; (14) SVL in females 23.4–28.2 mm (n = 7), in males 17.5–21.1 mm (n = 7).

No other species of Phrynopus has such a combination of dorsal folds and masked dark face. Among Phrynopus species lacking a tympanic annulus (present in P. auriculatus , P. montium , and P. peruanus ) and the dentigerous processes of vomers (character status for P. personatus in parentheses), P. capitalis is much larger, lacks dorsolateral folds and has a different dorsal coloration pattern, possesses bulbous digital tips (rounded) and wears lateral fringes in toes and fingers (absent). Phrynopus dumicola does not have a distinctive face mask, has smaller nasal bones, a uniform dorsal coloration, and no light spots on groin whereas P. valquii has a weakly defined canthus rostralis (sharp), much lighter coloration and bulbous digits (rounded in personatus ). Among species that exhibit bright coloration on groin ( P. badius , P. bracki , P. dagmarae , P. heimorum , P. interstinctus , P. nicoleae , P. paucari , P. peruanus , and P. vestigiatus ), P. peruanus has a tympanic annulus (absent); P. bracki has dentigerous processes of vomers (absent), and P. badius has smooth uniform brown ventral skin, scattered with minute white dots and labial bar and very prominent subarticular at the base of fingers (absent in P. personatus ); P. paucari has a bright yellow venter. Phrynopus vestigiatus shares with P. personatus the presence of X ridges on the scapular region, but has smooth dorsal skin, is smaller with single known female having SVL= 18.8 mm (28.2 mm), has minute dentigerous processes of vomers (absent), and a dark venter with small white flecks (light flecks absent). Phrynopus dagmarae possesses light groin marks sharply bordered by black, but differs from P. personatus by having the dentigerous processes of vomers and labial bars (both absent), different dorsal and ventral coloration, and bulbous tip of fingers (rounded, not expanded). Phrynopus horstpauli has dorsal markings and large nasal bones like P. personatus , but differs by its larger size, by lacking bright coloration on groin, and by having Tshaped terminal phalanges (knobbed) and yellow-copper iris (upper iris golden with brown reticulations). Phrynopus miroslawae is similar to P. personatus in having a supratympanic fold and broad dark canthal and postorbital stripes delineating a face mask, and by bearing dorsolateral and occipital folds, but differs by lacking bright coloration on groin, and by having bulbous digital tips (round), areolated ventral skin, and dorsal gray coloration with dark blotches. Phrynopus nicoleae is similar to P. personatus in having first finger shorter than second, and in bearing X-shaped dorsal marks and occipital and dorsolateral folds, but differs by its longer snout, and by having labial bars and vomerine teeth. Among species of Phrynopus lacking tympanum and dentigerous processes of vomers, P. curator also has dorsal Y shaped ridges, and tubercles, but has a weakly areolate ventral skin (aerolate), three tubercles on upper eyelid (dermal ridge with low tubercles), a conical tubercle on heel (absent), longer limbs (TL/SVL= 49% in P. curator , 33-35 % in P. personatus ) and lacks bright coloration in the groin (present in P. personatus ). Species of the Pristimantis orestes group bearing reduced digital pads may externally resemble Phrynopus personatus ; those are P. atrabracus , P. chimu , P. cordovae , P mariaelenae , P. melanogaster , P. pataikos , P. pinguis , P. seorsus , P. simonsii , P. stictoboubonus , P. stipa ; all of which, except P. simonsii , have either a visible tympanum, or the tympanic annulus is weakly defined or concealed beneath the skin (in P. melanogaster , P. pataikos , P. stictoboubonus , P. seorsus and P. stipa ). Furthermore, P. stipa , has a dark dorsal skin, but differs from Phrynopus personatus by lacking medial dorsal ridges and dorsolateral folds, and bearing prominent dentigerous processes of vomers (absent in P. personatus ). Pristimantis simonsii also lacks the dentigerous processes of vomers, but lacks dorsal folds and a distinctive face (both present in P. personatus ), and bright yellow or white spots on groin, on black ground (present). Pristimantis attenbouroughi differs by bearing dentigerous processes of vomers (absent), by having the upper eyelid without dermal ridge (dermal ridge present, bearing some low rounded tubercles), areolate ventral skin (finely areolate), heel with a small conical ulnar and tarsal tubercles (absent), and a short cloacal sheath (absent).

Description of holotype. Head narrower than body, longer than width; HW 95% of HL; HW 37.8% of SVL; interorbital area flat, no cranial crests; upper eyelid bearing several round supraorbital tubercles, EW 57% of IOD,; snout semicircular in dorsal view, slightly truncate in profile; snout short, E-N 69% of eye length; canthus rostralis sharp in lateral view, loreal region strongly sloping outward to lip, slightly concave; nostrils weakly protuberant, directed dorsolaterally; lips not flared; tongue large, oval, posterior one-fourth free; choanae small, round, situated well anterolaterally on palate, fairly separated; dentigerous processes of vomers absent; tympanic annulus and tympanic membrane absent, supratympanic fold thick and prominent; several postrictal tubercles present, extending onto the loreal area.

Skin of dorsum shagreened; prominent but incomplete dorsolateral ridges; flanks and upper surfaces of limbs bearing scattered conical warts, lateral surfaces of limbs and paravertebral area bearing low small tubercles; an Xshaped middorsal ridge formed by a series of closely packed warts starting postorbitally and joining the middorsal line at the middle of the back, shorter and less prominent on posterior half; skin on throat shagreened, that of venter finely areolate; an outer row of tubercles present on forearms; palmar tubercle ovoid, thenar rounded barely defined; subarticular tubercles low, simple; finger tips not bulbous; first finger shorter than second.

Tibia length 33% of SVL; showing an external row of tubercles on the heel and tarsal area, as an extension of the warts on lateral surfaces of limbs; outer metatarsal tubercle flat, oval, barely defined, inner slightly smaller than outer, rounded; supernumerary and subarticular tubercles not evident. Toes lacking lateral fringes and basal webbing.

Color in life. The dorsal ground color was brown to reddish brown; venter, throat and inner forelimbs were brown-red to red; bright to light rounded yellow spots on groin sharply bordered with black. Face and dorsal surfaces as in preservative. Iris golden with fine dark reticulations on upper half, lower half dark brown.

TABLE 1. Measurements and proportions of two new species of Phrynopus . For measurements abbreviations see Material and Methods. Mean and standard deviation are followed by range. P. dumicola sp. nov. P. personatus sp. nov.

♂ (n=6) ♀ (n=5) ♂ (n=7) ♀ (n=7) SVL 20.1 + 0.5 24.1 + 0.6 19.2 + 0.5 25.3 + 0.6 (18.7–22.6) (22.3–25.3) (17.5–21.1) (23.4–28.2) HL 6.9 + 0.2 8.0 + 0.4 6.8 + 0.2 8.9 + 0.3 (6.2–7.7) (7.2–9.4) (6.0–7.4) (7.7–10.0) HW 7.5 + 0.2 8.6 + 0.4 7.0 + 0.2 9.3 + 0.3 (6.6–8.0) (7.9–10.1) (6.5–7.6) (8.1–10.4) THL 7.2 + 0.1 8.8 + 0.4 7.8 + 0.2 9.3 + 0.2 (6.9–7.5) (8.1–9.5) (7.2–8.5) (8.7–10.2) TL 6.9 + 0.1 8.2 + 0.2 6.7 + 0.2 8.4 + 0.1 (6.5–7.1) (7.6–8.9) (6.1–7.7) (8.2–9.2) FL 8.5 + 0.2 10.0 + 0.2 8.5 + 0.3 10.0 + 0.3 (7.9–9.5) (9.5–10.8) (7.0–9.4) (8.3–11.3) IOD 2.3 + 0.1 2.6 + 0.1 2.2 + 0.1 2.8 + 0.1 (2.1–2.7) (2.2–2.9) (1.9–2.5) (2.4–3.3) EW 1.8 + 0.1 2.2 + 0.1 1.5 + 0.1 1.9 + 0.2 (1.6–2.1) (1.9–2.5) (1.3–1.9) (1.4–2.8) E-N 1.5 + 0.1 1.9 + 0.2 1.6 + 0.2 1.8 + 0.1 (1.4–1.6) (1.3–2.7) (1.3–2.4) (1.4–2.4) IND 2.0 + 0.1 2.6 + 0.2 2.0 + 0.2 2.6 + 0.1 (1.8–2.2) (2.2–3.1) (1.4–2.7) (2.2–3.1) ED 2.5 + 0.2 3.1 + 0.2 2.4 + 0.2 3.0 + 0.3 (2.0–2.9) (2.6–3.8) (2.0–3.2) (2.3–3.9)

TL/SVL 34.9% 33.4%

HW/SVL 36.7% 36.6%

Color in preservative. The face is uniformly dark brown, with a faint line bordering the canthal stripe and the tympanic fold. The tympanic and loreal regions underneath the faint line are dark brown, and form a pattern resembling a mask. The upper lip is bordered by small cream blotches. A thin, pale cream middorsal stripe extends from the internasal area to the vent. Above the cloacal opening, the stripe divides and extends across the thighs to the lower edge of the knee, continuing on the posterior side of shanks to reach the heel, and forming an irregular broad cream stripe (yellow in life) on the inner surface of the shank. Dorsal coloration is gray-brown above, with dorsal folds, warts and ridges bordered with faint and dark brown, as on the face. Venter and concealed surfaces of limbs, brown with dirty cream blotches. Rear surfaces of thigh and knees are dark brown.

Measurements (in mm) of the holotype. SVL 24.5, TL 8.1, FL 9.7, HW 9.3, HL 9.7, IOD 2.7, EW 1.6, IND 2.3, ED 2.3, E-N 1.6.

Variation. Tibia length is 33% of SVL in females (n=7) and 35% in males (n=7). In some specimens, the postympanic ridge does not continue over the eye. Some specimens bear small oblique ridges, formed by a row of round tubercles (MUSM 3811 and MUSM 381, AMNH 134153), in the interorbital area. The dorsolateral folds are not continuous in MUSM 33100 and completely absent in MUSM 33091. This last specimen has an additional white spot (yellow in life) in the inner thigh, and two in the inner shank, as does MUSM 3811. In life, MUSM 9073, 90–74 and 9076, had very dark dorsal coloration and some ventral irregular spots. The thin pale cream middorsal line is persistent in all specimens but MUSM 33100, where the line is ill defined.

Etymology. The specific name personatus is a Latin adjective meaning masked and is used in reference to the distinctive face mask of this species.

Distribution and ecology. Phrynopus personatus is known only from a narrow elevational range around the type locality in the upper reaches of the continuous mountain forests of the Abiseo National Park. Fourteen specimens were found in wet leaf litter of flat areas, all in the narrow elevational range from 2890 to 3110 m near El Mirador (3000 m). The area is a continuous tropical montane rain forest about 15 m tall. Arboreal ferns are common and tree trunks and ground surfaces are covered with moss, liverworts and lichens. Individuals of P. personatus were notably inactive before and after capture. The reproductive stage of the holotype indicates that the species may reproduce in the dry season. Males weighed 0.65– 0.95 g, females 1.90–3.05 g. Although no eggs were found with the specimens, the large size of the well-developed eggs in the holotype strongly suggests direct development. Some specimens had trombiculid mites under the skin (see cysts in Figures 13 B, F).

Figures

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.

FIGURE 13. Live specimens of Phrynopus personatus sp. nov. (A, B) MUSM 33121, (C, D) MUSM 33100 and (E, F) MUSM 33091 from near El Mirador, 2890 m, Río Abiseo National Park, San Martín, Peru. Photographs by A. Catenazzi.