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

neighboring favorable habitats. Almost half of this area is inside the Parque Nacional Podocar- pus, basically delimiting the park’s northern border. Parque Nacional Podocarpus was declared officially in 1982 and comprises an area of 146,289 ha. The national park has a very irregular topography covering altitudes from 900 to 3650 m, with large areas of diverse natural habitats [17] and it is an important part of the Tropical Andes hotspots, the richest in biodiver- sity on the planet [18, 19]. The study site has an altitudinal range between 1700 and 3000 m a.s. l. and consists of evergreen upper montane forest (on the western slope, near Loja, up to 2700 m), subpáramo or evergreen elfin forest (on the crest, between 2700 and 3000 m), evergreen upper montane forest (on the eastern slope, between 2700 and 2100 m), and evergreen lower montane forest (on the eastern slope, near El Tambo, bellow 2100 m [20]) (Fig 2). In the study area, the average annual precipitation is around 4865 mm [21] and the average annual temper- ature around 7.3 ˚C [22]. Fieldwork was carried out between January 2016 and March 2020 (a total of 97 days) in Abra de Zamora and in several neighboring areas: Huacapamba (about 10 km to the north; 3.9061˚ S, 79.1909˚ W; datum WGS84), Reserva Madrigal del Podocarpus (about 9 km to the south; 4.0595˚ S, 79.1632˚ W) and the Cajanuma sector of Parque Nacional Podocarpus (about 13 km south to Abra de Zamora; 4.1163˚ S, 79.1719˚ W) (Fig 1). Additional specimens were collected from the wetland complex of Oña, Nabo´n, Saraguro and Yacuambi (3.5810˚ S,

79.0939˚ W), Bosque Protector Washapamba (3.6615˚ S, 79.2738˚ W), Ramos Urcu (at about 5 km west to San Lucas; 3.7111˚ S, 79.3019˚), Reserva Tapichalaca (4.4859˚ S, 79.1477˚ W), and Reserva Cerro Plateado (4.6565˚ S, 78.8699˚ W) in Loja and Zamora Chinchipe provinces. Fig 2. Abra de Zamora. A. General view of the higher part of Abra de Zamora (2800 m a.s.l.) with subpa´ramo ecosystem in the foreground. These terrestrial bromeliads are the type of microhabitat where Gastrotheca psychrophila specimens were collected in the ‘60s and ’70s. In the background the old road that connected Loja with Zamora (Vı´a Antigua Loja-Zamora) is visible. B. Subpa´ramo ecosystem of the higher part (2750 m a.s.l.), habitat of Pristimantis balionotus, P. colodactylus, P. matildae sp. nov., P. samaniegoi sp. nov., and P. versicolor. C. Evergreen upper montane forest ecosystem (2600 m a.s.l.), habitat of P. andinognomus, P. atratus, P. cryptomelas, P. samaniegoi sp. nov., and P. versicolor. D. Evergreen lower montane forest ecosystem (1850 m a.s.l.), habitat of Bolitoglossa sp., Rhinella margaritifera, Gastrotheca testudinea, P. galdi and an undescribed Pristimantis species. https://doi.org/10.1371/journal.pone.0238306.g002 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 4 / 59