Three new species of Pristimantis (Amphibia, Anura, Craugastoridae) from upper montane forests and high Andean grasslands of the Pui Pui Protected Forest in central Peru

We describe three new species of Pristimantis from the upper montane forests and high Andean grasslands of the Pui Pui Protected Forest and its close surroundings (Region Junin, central Peru) and compare them morphologically and genetically with other taxonomically and biogeographically relevant species of Pristimantis. All three new species have the skin on dorsum shagreen with scattered tubercles, discontinuous dorsolateral folds, tuberculate flanks, and the skin on venter areolate. Pristimantis bounides sp. nov. is known from two localities outside the Pui Pui Protected Forest in upper montane forests between 3350 and 3463 ma.s.l. and is characterized by a snout-vent length of 18.2-21.0 mm in males (n = 3), and 21.6-24.4 mm in females (n = 4), by having a tympanum, males with vocal slits, and discs of digits slightly expanded with circumferential grooves. In life, dorsal and lateral ground coloration is pale grayish brown, orange brown, yellowish brown or reddish brown with dark grayish-brown marmorations, and a pale gray, pale greenish gray or creamish white venter with or without dark gray mottling. Pristimantis humboldti sp. nov. is known from one locality inside the Pui Pui Protected Forest, in upper montane forest at 3318 ma.s.l., and is characterized by a snout-vent length of 17.2-20.6 mm in males (n = 3), and 19.7-25.7 mm in females (n = 6), by having a tympanum, males with vocal slits, and discs of digits expanded with circumferential grooves. In life, dorsal and lateral ground coloration is orange brown with brownisholive blotches, orange brown with grayish-brown blotches and flecks, reddish brown with grayish-brown blotches or grayish brown with orange brown blotches; throat, chest, belly, anterior and ventral surfaces of thighs, tibia, and axilla are dark gray and pale gray mottled with white and pale gray spots of different sizes and density. Pristimantis puipui sp. nov. is known from one locality inside the Pui Pui Protected Forest, in the puna at 3890 ma.s.l., and is characterized by a snoutvent length of 16.1-17.1 mm in males (n = 3), and 20.6-22.4 mm in females (n = 4), by lacking a tympanum, lacking males with vocal slits, and tips of digits narrow without circumferential grooves. In life, dorsal and lateral ground coloration is pale orange brown, reddish brown or grayish brown with or without grayish-brown mottling, and the venter is pale cream and pale gray mottled. A molecular phylogenetic analysis based on mitochondrial and nuclear DNA sequences inferred that the three new species belong to the Pristimantis danae species Group distributed in the montane forests and high Andean grasslands of central Peru, including P. albertus, P. aniptopalmatus, P. ornatus, and P. stictogaster. With the three new species, 133 species of Pristimantis are currently known from Peru, eight of which inhabit the puna.