Fifty years after: A taxonomic revision of the amphibian species from the Ecuadorian biodiversity hotspot Abra de Zamora, with description of two new Pristimantis species

Diagnosis. Pristimantis versicolor is a medium sized species, distinguished by the following combination of traits: (1) skin on dorsum shagreen with fine tubercles; skin on venter areolate; discoidal fold visible, thoracic fold prominent; dorsolateral folds absent; (2) tympanic mem- brane and tympanic annulus prominent, its length about 45% of the length of eye; supratympa- nic fold present; (3) snout subacuminate in dorsal view, rounded in profile; canthus rostralis angular, straight or slightly concave in dorsal view, angular in profile; (4) upper eyelid usually bearing several larger tubercles (trait more visible in life), about 85% IOD in females and 97% IOD in males; cranial crests absent; (5) dentigerous processes of vomers prominent, oblique, ovoid, separated medially by distance equal to twice than the width of processes; each pro- cesses bearing 3 to 4 teeth; (6) males lacking vocal sac and slits; nuptial pads present; (7) Finger I shorter than Finger II; discs on fingers broadly expanded, elliptical; circumferential grooves present; fingers bearing weak lateral fringes or none; (8) subarticular tubercles prominent, round; supernumerary palmar tubercles evident; palmar tubercle partially divided into a larger (inner) and a smaller (outer) tubercles; thenar tubercle oval, equal in size with the inner palmar tubercle; (9) ulnar tubercles low, diffuse; (10) heel with one large and several smaller tubercles (trait more visible in life); outer edge of tarsus bearing a row of small, conical tubercles; inner

tarsal tubercles coalesced into a short tarsal fold; (11) inner metatarsal tubercle ovoid, about 3 to 4x round, conical (in profile), outer metatarsal tubercle; supernumerary plantar tubercles present, conical; (12) Toe V much longer than Toe III; discs on toes broadly expanded, ellipti- cal, slightly smaller than those of fingers; circumferential grooves present; toes lacking lateral fringes; webbing absent; (13) in life, dorsum brown or green, with dark markings (chevrons, interorbital bar, canthal and supratympanic stripes); flanks with vertical bars or spotted; limbs usually with transverse bars; venter cream with black or dark brown reticulations; no markings in axilla, groin or on concealed limb surfaces; iris whitish bronze with a broad reddish median horizontal streak, that almost covers all the lower part of the iris and fine black reticulations; (14) SVL 29.4–32.2 mm in adult females (30.8 ± 1.40 SD, n = 3) and 22.9–25.4 mm in adult males (24.0 ± 1.08 SD, n = 4). Variation. Morphometric variation is shown in Table 3. The “versicolor” (variegated) name is very fitting because this species displays an incredible color variation. Many individu- als have a general brown coloration with green blotches and whitish markings like female MUTPL 390 (Fig 27A) or a general green coloration (MUTPL 313, Fig 27F). In some Fig 27. Color variation of Pristimantis versicolor in life. A. female: MUTPL 390; B–C males: B. MUTPL 112, C. MUTPL 740; D–E subadults: D. MUTPL 497, E. MUTPL 293; F. juvenile: MUTPL 313. Abra de Zamora (A, B, D, E,

F), Reserva Tapichalaca (C). Most common coloration of this species is presented in A and F. https://doi.org/10.1371/journal.pone.0238306.g027 PLOS ONE Taxonomic revision of the amphibian species from Abra de Zamora PLOS ONE | https://doi.org/10.1371/journal.pone.0238306 September 10, 2020 48 / 59