(Figs 1–3, Table 2). CORBIDI 18226, an adult male (Figs 2, 3) from 13°22'12.14''S; 71°6'49.82''W (WGS84), 2795–2850 m a.s.l., Quispillomayo valley, Área de Conservación Privada (ACP) Ukumari Llaqta, Distrito Paucartambo, Provincia Paucartambo, Departamento de Cusco, Peru, collected by A. Catenazzi, A. Shepack, M. I. Diaz and A. Ttito on 27 May 2016.

(Fig. 4, Table 2). Four specimens: two females, CORBIDI 18224 and MUBI 14654, and one male, CORBIDI 18225, collected with the holotype on 27 May 2016; and one male, MUBI 14655 collected on 28 May 2016.

Three juveniles, CORBIDI 18227, 18228, and MUBI 14665, collected with the holotype and paratopotypes on 27 May 2016.

A new species of Bryophryne as defined by Duellman and Lehr (2009), Hedges et al. (2008), and Padial et al. (2014). Frogs of the genus Bryophryne are morphologically similar and closely related to Barycholos, Holoaden, Noblella and Psychrophrynella (Chaparro et al. 2015; Hedges et al. 2008; Heinicke et al. 2007; Padial et al. 2014). Genetic data confirm generic placement of the new species within Bryophryne (Table 1). We found substantial genetic distances (uncorrected p-distances from 3.7–6.7%; Table 1) between B. phuyuhampatu and congeneric species for which mitochondrial sequence data were available (B. bakersfield, B. bustamantei, and B. cophites). The most closely related species is B. bakersfield (16S uncorrected p-distance: 3.7–4.1%), followed by B. cophites (5.6–6.2%) and B. bustamantei (5.6–6.7%). Regarding species from other genera, B. phuyuhampatu had genetic distances ranging from 12.4% (Psychrophrynella guillei) to 20.8% (Barycholos pulcher). In addition to the molecular data, the new species is assigned to Bryophryne rather than any of the other genera on the basis of overall morphological resemblance with the type species B. cophites, including head narrower than body, short limbs, and tympanic membrane and annulus usually absent (absent in most species of Bryophryne, except for B. flammiventris and B. gymnotis), and geographic distribution within the Departamento Cusco, where all other species of Bryophryne occur.

A new species of Bryophryne characterized by: (1) skin on dorsum shagreen; skin on venter areolate, discoidal fold absent, thoracic fold present; dorsolateral folds irregular and discontinuous; (2) tympanic membrane and tympanic annulus absent; (3) snout rounded in dorsal view and in profile; (4) upper eyelid with two small tubercles, narrower than IOD; cranial crests absent; (5) dentigerous process of vomers absent; (6) vocal sac and slits absent; nuptial pads absent; (7) Finger I much shorter than Finger II; tips of digits slightly pointed; (8) fingers lacking lateral fringes; (9) outer edge of forearm bearing small tubercles; (10) heel bearing minute tubercles; inner tarsal fold absent; outer edge of tarsus bearing small tubercles; (11) inner metatarsal tubercle prominent, ovoid, of similar relief and slightly larger than ovoid, outer metatarsal tubercle; supernumerary plantar tubercles indistinct; (12) toes lacking lateral fringes; webbing absent; toes III and V about equal in length; tips of digits slightly pointed; (13) in life, dorsum tan to green and brown with dark brown markings, greenish blue on lower flanks; some specimens with a yellow middorsal line extending from tip of snout to cloaca and to the posterior surface of thighs; interorbital bar present; chest, belly and ventral parts of forearms and legs dark brown with grayish blue flecks; throat brown with flecks turning from gray-blue to copper near tip of mouth; palmar and plantar surfaces brown with lighter fingers and toes; (14) SVL 14.2–16.9 in males (n = 3), 22.2–22.6 in females (n = 2).

The new species differs from other members of the genus by having green coloration on dorsum and blue coloration on flanks and ventral parts. Furthermore, B. phuyuhampatu differs from other species by the following combination of characters (condition for comparing species in parenthesis): from B. abramalage by having proportionally longer feet with FL/SVL from 0.41–0.45 (0.37–0.42), narrower head with HW/HL from 0.85–0.86 (0.97–1.07), and inner metatarsal tubercle larger than outer metatarsal (inner half the size of outer metatarsal tubercle); B. flammiventris and B. gymnotis by lacking a tympanum (present), from B. bakersfield and B. bustamantei by having discontinuous dorsolateral folds (continuous), from B. cophites by females being much smaller (22.6 mm vs. 35.8 mm), from B. hanssaueri by lacking bright orange coloration on throat (present), from B. nubilosus by having toes III and V similar in length (toe V > III), and from B. zonalis by having blue-gray mottled coloration on belly (distinctive black mottled coloration, variably confined to lower portion of belly). The new species further differs from B. gymnotis by having vomers lacking dentigerous processes (present), from B. cophites by males lacking nuptial pads (present), and from B. bakersfield, B. bustamantei, B. flammiventris and B. gymnotis for males lacking vocal slits (present).

Bryophryne phuyuhampatu (max. SVL 22.6 mm) is much smaller than B. bakersfield (31.1 mm), B. cophites (35.8 mm; pers. obs.), B. hanssaueri (29.3 mm; pers. obs.), and B. zonalis (32.4 mm), smaller than B. bustamantei (23.4 mm), B. flammiventris (24.1), B. hanssaueri (24.6 mm), and about the same size of B. abramalagae (20.1 mm) and B. nubilosus (26.0 mm; pers. obs.). Five other small species of craugastorid frogs of the subfamily Holoadeninae are known to occur in montane forests and high Andean grasslands south of the Apurimac canyon in Peru: Noblella madreselva, N. pygmaea, Psychrophrynella bagrecito, P. chirihampatu and P. usurpator, which all possess a visible tympanic annulus.

Adult male (16.9 mm SVL); head narrower than body, its length 33% of SVL; head wider than long, head length 83% of head width; head width 32% of SVL; snout short, rounded in dorsal and lateral views (Fig. 2), eye diameter 35% of head length, its diameter 1.2 times as large as its distance from the nostril; nostrils slightly protuberant, close to snout, directed dorsolaterally; canthus rostralis slightly straight in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; upper eyelids with two small tubercles; upper eyelid width 46% of interorbital distance; interorbital region flat, lacking cranial crests; eye-nostril distance 81% of eye diameter; supratympanic fold short and weak; tympanic membrane and tympanic annulus absent; postrictal tubercles absent. Vocal sac and vocal slits absent. Choanae ovoid, small, positioned far anteriorly and laterally, widely separated from each other; dentigerous processes of vomer and vomerine teeth absent; tongue large, ovoid, about 2.5 times as long as wide, not notched posteriorly.

Skin on dorsum shagreen with small, scattered tubercles; dorsolateral folds discontinuous, extending from posterior margin of upper eyelid to sacral region; skin on flanks tuberculate; skin on throat smooth, skin on chest, and belly areolate; thoracic fold present, discoidal fold absent; cloaca slightly protuberant, cloacal sheath short, cloacal region without tubercles. Outer surface of forearm with minute tubercles; palmar tubercle flat and oval, approximately same length but twice the width of elongate, thenar tubercle; few supernumerary tubercles low, ovoid; subarticular tubercles prominent, ovoid in ventral view, rounded in lateral view, largest at base of fingers; fingers lacking lateral fringes; Finger I much shorter than Finger II; relative lengths of fingers 3 > 4 = 2 > 1 (Fig. 3); tips of digits slightly pointed, lacking circumferential grooves (Fig. 3); forearm lacking tubercles.

Hindlimbs short and robust, tibia length 38% of SVL; foot length 39% of SVL; upper surfaces of hindlimbs shagreen with scattered, minute tubercles; posterior surface of thighs tuberculate to areolate, ventral surface areolate; heel with minute tubercles (not visible in preservative); inner edge of tarsus without tubercles, outer edge of tarsus with small tubercles; inner metatarsal tubercle prominent, ovoid, of similar relief and slightly larger than ovoid, outer metatarsal tubercle; supernumerary plantar tubercles indistinct; subarticular tubercles low, ovoid in dorsal view; toes lacking lateral fringes, not webbed; toe tips weakly pointed, not expanded laterally, about as large as those on fingers; relative lengths of toes: 4 > 3 = 5 > 2 > 1 (Fig. 3); foot length 32% of SVL.

Measurements of holotype (all in mm): SVL 16.9, TL 6.5, FL 6.6, HL 5.5, HW 5.4, ED 1.6, IOD 2.4, EW 1.1, IND 1.5, E–N 1.3.

(Fig. 2). Dorsum green and brown with a dark brown marking extending from the interorbital bar to a mid-dorsal longitudinal band, a horizontal dark mark near the sacral region, and an oblique dark band on each flank. Dorsal surfaces of arms and legs dark brown, with transverse dark bars on forearms and hind limbs. Lower flanks with greenish blue flecks. Chest, belly and ventral parts of forearms and legs dark brown with grayish blue flecks. Iris grayish blue with a medial copper band. Throat brown with flecks turning from gray-blue to copper near the tip of the mouth. Palmar and plantar surfaces brown; tips of fingers and toes light brown to yellow.

(Fig. 2). Similar to coloration in life, but dorsal surfaces grayish tan with higher contrast of dorsal markings. Ventral surfaces beige to brown with cream flecks.

Coloration in life is based on field notes and photographs taken by A. Catenazzi of the paratopotypes (Fig. 4; photographs available through Calphoto database). The amount of dorsal green coloration varies among specimens. While juvenile MUBI 14665 and male MUBI 14655 are similar to the holotype in having a generally greenish dorsum, all other specimens have dark tan to brown dorsum, with just a few tubercles colored green. Female MUBI 14654, male CORBIDI 18225 and juvenile CORBIDI 18228 have a yellow middorsal line extending from the tip of the snout to the cloaca and to the posterior surface of the thighs.

The summary of measurements of all types is reported in Table 2.

The specific name phuyuhampatu is a combination of Quechua words used in apposition meaning “toad” (“hampa'tu”) that lives in the “fog” (“phuyu”).

Bryophryne phuyuhampatu was discovered during a rapid amphibian survey in the upper Quispillomayo Valley (Fig. 5A) from 22 to 31 May 2016. The Quispillomayo torrent (Fig. 5B) is a tributary of the Nusiniscato River, which reaches the Araza River downstream of Quincemil, in the upper Madre de Dios drainage. During the inventory high-Andean grasslands (puna; 3350–4515 m a.s.l.), a forest patch of tasta (Escallonia myrtillioides), kishuar (Buddleja incana) and qeñua (Polylepis incana) at 4280 m a.s.l., montane scrub, disturbed areas and other transitional formations along the treeline around 3350 m a.s.l., and the montane cloud forest from 2780–3350 m a.s.l. were sampled. Frogs were searched for under rocks, logs, mosses, and in the leaf litter and the understory in the montane forest. All but one specimens of B. phuyuhampatu were found under mosses in the cloud forest around 2850 m a.s.l. (Fig. 5C). Male MUBI 14655 was found ~250 m from this site, under rocks and mosses under the riparian vegetation at the confluence of a small stream at 2795 m a.s.l. Two sympatric frogs, Gastrotheca cf. excubitor and Psychrophrynella chirihampatu, were found under rocks in disturbed habitats (i.e., along streams, landslides) but not in the cloud forest. Two additional amphibian species, Bryophryne sp. and B. cf. zonalis, were found along with G. cf. excubitor in the grasslands from 3100–3650 m a.s.l.

Both female paratopotypes had large eggs in their ovaries, indicative of terrestrial breeding and direct development: CORBIDI 18224 contained 15 eggs averaging 1.58 ± 0.05 mm in diameter (range 1.20–1.80 mm), while MUBI 16654 contained 16 eggs averaging 2.44 ± 0.03 mm in diameter (range 2.30–2.60 mm).

The type locality (and known distribution range) of the new species lies within the Área de Conservación Privada Ukumari Llaqta (Catenazzi and Ttito 2016), a protected area recognized by Peruvian environmental ministerial decree N° 301–2011-MINAM in December 2011. The upper puna and transitional habitats, as well as a narrow elevational band around the treeline are used for agriculture (potato cultivation), livestock (llamas grazing), fishing (exotic trout), and timber extraction. These land use patterns appear sustainable, and the grasslands at Patawasi (3350–3450 m a.s.l.) are in excellent conditions, with large bunchgrasses supporting large populations of Bryophryne sp. and B. cf. zonalis. There is little indication of human disturbance in the cloud forest, and the main use seems to be limited to trout fishing.