Phrynopus sancristobali

Description Phrynopus sancristobali sp. nov. is distinguished by a combination of the following characters: (1) skin on dorsum and flanks with large, round and irregular pustules, forming irregular ridges; skin on the belly strongly areolate; discoidal fold absent, thoracic fold absent; dorsolateral folds absent; postocular fold prominent; (2) tympanic membrane and tympanic annulus absent; (3) snout short, round in dorsal view, curved antero-ventrally in lateral view; (4) upper eyelid without tubercles; width of upper eyelid slightly narrower than IOD, cranial ridges absent; (5) dentigerous processes of vomers absent; (6) vocal slits and nuptial pads absent; (7) Finger I shorter than Finger II; rounded and bulbous fingertips, lacking discs or grooves; (8) fingers without basal webbing and without lateral fringes; (9) ulnar tubercles absent, tarsal tubercle absent; (10) thenar and palmar tubercles small, internal tarsal fold absent; (11) internal metatarsal tubercle rounded, larger than external, ovoid metatarsal tubercle; subarticular tubercles entire on all fingers, supernumerary plantar tubercles absent; (12) toes without lateral fringes, basal webbing absent, Toe V slightly longer than Toe III, bulbous fingertips, no discs; (13) in life, dorsum and dorsal parts of extremities light brown, with dark brown reticulations that appear to divide the pustules; flanks golden brown with gray reticulations and dark brown; palms of hands orange, Toes IV and V dark brown, plantar surface of foot of a more intense orange color; belly cream to yellow with gray to light brown crosslinks, groin and throat orange; iris silvery with dark brown marks; (14) SVL 20.7 and 22.2 mm in two females and 19.5 in one male.

Description of holotype Head nearly as wide as body, wider than long; head width 107 % of head length; head width 37 % of snout-vent length; head length 35 % of SVL. Snout short, rounded in dorsal view, curved anteroventrally in lateral view (Figs. 3 – 5), eye-to-nostril distance 48 % of eye diameter; nostrils not protuberant; canthus rostralis slightly curved in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; upper eyelid lacking tubercles, narrower than interorbital distance (EW 66 % of IOD); postocular and tympanic region with prominent folds (Fig. 3); tympanic membrane and tympanic annulus absent, tympanic region with discontinuous pustules; choanae small, ovoid, partly concealed by palatal of maxilla, dentigerous processes of vomers absent; tongue broad, about 1.7 as long as wide, notched posteriorly, posterior half free; vocal slits absent. Skin on dorsum with circular pustules of varying dimensions (Fig. 4); skin on flanks shagreened with scattered rounded pustules; skin on throat, chest and belly strongly areolate (Fig. 3 B); discoidal and thoracic folds absent; cloacal sheath absent; cloacal region with prominent pustules. Outer surface of forearm with prominent pustules; outer palmar tubercle barely visible, low, ovoid, smaller than ovoid inner palmar tubercle; supernumerary tubercles absent; subarticular tubercles absent; fingers without lateral fringes; Finger I slightly shorter than Finger II (Fig. 5); tips of digits rounded, bulbous, lacking circumferential grooves. Hind limbs short and slender, tibia length 34 % of SVL; foot length 37 % of SVL; dorsal surface of hind limbs with irregular pustules; anterior surfaces of thighs smooth, posterior surfaces of thighs pustulate; heel with low rounded, irregular tubercles; outer surface of tarsus bearing pustules like those on dorsum; outer metatarsal tubercle rounded and prominent, slightly larger than the ovoid inner metatarsal tubercle; supernumerary plantar tubercles absent; subarticular tubercles low, not distinguishable; toes without lateral fringes; basal webbing absent; toe tips bulbous, rounded, lacking circumferential grooves, about as large as those on fingers; relative lengths of toes: 1 <2 <3 <5 <4; Toe V slightly longer than Toe III (Fig. 5). Measurements of holotype (in mm). SVL, 19.5; TL, 6.6; HL, 6.8; IND, 1.7; IOD, 2.3; EW, 1.6; HW, 7.3; FL, 7.3; E-N, 1.1; ED, 2.2 (Table 1). Variation Female PFAUNA 550 is larger (SVL = 22.2 mm) than both the holotype and male PFAUNA 551 (SVL = 20.7 mm; Table 1). Furthermore, the female has the first subarticular tubercle entire in Finger I and IV, and divided in Fingers II and III (all subarticular tubercles entire in both males; Fig. 7). Coloration of holotype in life The holotype had a light brown to golden dorsum, with dark brown reticulations that appear to divide the pustules, and similar coloration on the dorsal part of the limbs; the flanks were golden brown with gray and dark brown reticulations; the belly was cream to yellow with gray to light brown reticulations, the groin and throat were orange; there is a dark brown halo on the nose and on the eye, extending as a post ocular stripe that ends in the tympanic palms and soles were orange, but fingers IV and V were dark brown in dorsal and ventral view, fading to orange at about half of palms (Fig. 3). Coloration of holotype in preservative The holotype in preservation maintains the same general color patterns described above, only decreasing in intensity (Fig. 4).

Figures 3 – 7, Table 1 urn: lsid: zoobank. org: act: 9 B 5329 CC- 7045 - 4 EE 5 - B 71 F-A 6 DD 3 ACC 0 DFF

Diagnosis Phrynopus sancristobali differs from P. auriculatus, P. montium, P. mariellaleo and P. peruanus by lacking tympanic membrane and annulus; from P. auriculatus, P. remotum, P. inti, P. bracki, P. dagmarae, P. interstinctus, P. lechriorhynchus, P. kauneorum, P. kotosh, P. peruanus and P. vestigiatus by lacking dentigerous processes of vomers; from P. juninensis, P. kauneorum, P. inti, and P. apumantarum by the presence of prominent postocular folds; from P. chaparroi, P. thompsoni, P. paucari, P. heimorum, P. remotum, P. anancites, P. miroslawae, P. juninensis, P. kauneorum, P. tautzorum, and P. apumantarum by the presence of dorsal pustules with large, round and irregular warts; from P. remotum, P. anancites, P. miroslawae, P. juninensis, P. kauneorum, P. tautzorum, and P. apumantarum by the presence of pustules on dorsum; from P. inti, P. tribulosus, P. valquii, P. oblivius P. auriculatus, P. kauneorum, P. bracki, P. mariellaleo and P. tribulosus by having ventral skin strongly areolated. Phrynopus sancristobali most resembles P. bufoides, from which it differs by having irregular pustules on the dorsal part of the fore and hind limbs, maximum SVL 22.2 mm in females, ventral coloration in females cream with irregular yellow spots, and throat and ventral surface of arm and leg mostly orange (pustules arranged in longitudinal rows, maximum SVL 32.9 mm in females, venter and throat coloration in females gray with dark-brown spots irregularly distributed, and ventral surface of leg mostly dark-brown in P. bufoides).

Distribution and natural history Phrynopus sancristobali sp. nov. is only known from the type locality, which is characterized by the transition from humid puna to montane forest at elevations from 3796 to 3910 m a. s. l., in the upper watershed of the Caballuyoc river, tributary of the Mantaro river, within the Huanta province in the Ayacucho Department. The frogs were found under rocks in grasslands (Fig. 8) dominated by Stipa ichu, Stipa obtusa, Senecio cf. bukatii, Oreithales integrifolia, Halenia umbellata, Azorella creanata, and Chaptalia similis. Phrynopus sancristobali sp. nov. is sympatric with frog species Gastrotheca sp. and Pleurodema marmoratum, as well as the lizard Wilsonosaura josyi.

Etymology The specific epithet honors the Universidad Nacional San Cristóbal de Huamanga, alma mater of the first author, in recognition of its commitment to forming professional biologists with a holistic approach to biodiversity conservation.

Holotype PFAUNA 552, adult male, from Escalerapampa, District of Ayahuanco, Province of Huanta, Department of Ayacucho, Peru (12 ° 32 ’ 50 ” S, 74 ° 15 ’ 07 ” W, 3910 m a. s. l.) collected on 16 January 2020 by Vladimir Díaz and Randy Llantoy. Figures 3 – 5. Paratypes Two adults from near the type locality: female PFAUNA 550 (12 ° 33 ’ 8 ” S, 74 ° 15 ’ 02 ” W, 3910 m a. s. l.) and female PFAUNA 551 (12 ° 32 ’ 50 ” S, 74 ° 15 ’ 07 ” W, 3910 m a. s. l.) collected on 16 January 2020 by V. Díaz and R. Llantoy. Figure 6 – 7.