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

Remarks. This is an uncommon species in Abra de Zamora. Can be encountered on the eastern slope in evergreen upper montane forest (Fig 2C), between 2700 and 2500 m. Pristimantis galdi Jime´nez de la Espada, 1870 (Fig 29E) Common English name. Espada’s Robber Frog Common Spanish name. Cutı´n verde amazo´nico Identification. This small sized species is easily distinguished from all other Pristimantis species present in Abra de Zamora by the bright green dorsum, green iris, acuminate snout, flared lips, serrated cranial crests (especially in females), a conical tubercle on the upper eyelid and greatly expanded, truncate discs on Fingers II–IV [25]. Remarks. This is an uncommon species in Abra de Zamora. Can be encountered on the eastern slope in evergreen lower montane forest (Fig 2D), around 1900 m, near the El Tambo sector. Pristimantis lymani (Barbour and Noble, 1920) (Fig 29F) Common English name. Lyman’s Robber Frog Common Spanish name. Cutı´n de Lyman Identification. This species is easily distinguished from all the other Pristimantis species present in Abra de Zamora by its large size, prominent tympanum, smooth skin on the venter, reddish brown dorsum, canthal and postorbital dark stripes, chevrons on the back, and trans- verse bars on the limbs [25]. It is the only member of the Pristimantis conspicillatus species group, and the only terrestrial Pristimantis species from the area. Remarks. This is a common species in Abra de Zamora and the most common species in the nearby Loja city. Can be encountered on the western slope in evergreen upper montane

forest bellow 2600 m, and in the pasturelands neighboring the city. Additionally, 10 potential new species were encountered in Abra de Zamora: • from the western slope, in the vicinity of Loja city, Noblela aff. heyeri and two Pristimantis species (P. aff. cajamarcensis and P. aff. torresi); • from the eastern slope, heading to El Tambo, an undescribed species of salamander (Bolito- glossa sp.), Atelopus aff. halihelos, two undescribed species of glass frogs (Centrolene sp.), Noblela aff. heyeri, and three undescribed species of Pristimantis. Unfortunately, from the 29 species registered (belonging to six families, Plethodontidae, Bufonidae, Centrolenidae, Hemiphractidae, Strabomantidae and Telmatobiidae), one (A. aff. halihelos) was not encountered in the past ten years and three others (A. podocarpus, G. psy- chrophila, and T. cirrhacelis) were never seen again for over twenty years. Discussion Systematics Already in 1979, Lynch [9] observed the similarity between P. vidua and P. orestes and hypoth- esized a close relationship; this was later recognized as a distinct assemblage in 1997 by Lynch and Duellman [59], composed by three Ecuadorian species (P. orestes, P. simonbolivari and P. vidua). We now have the phylogenetic confirmation that indeed P. vidua is a member of the resurrected P. orestes group (sensu [29]), being closely related to the species of the P. orestes subgroup: P. andinognomus, P. orestes, P. cajanuma and an undescribed species (Fig 7). The situation was different for P. colodactylus, which, due to its particular morphological

features, was included in a different group. In the original description [9], Lynch hypothesized 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 53 / 59