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

of 3.56 notes/s (SD = 0.126) (S4 Table). The mean dominant frequency of the call was 2431.7 Hz (SD = 59.371), with a mean 90% bandwidth of 2331.8–2542.8 Hz (Table 2). The fundamen- tal frequency is not recognizable, but usually 4 harmonics are visible. Distribution. Pristimantis balionotus is known only from Abra de Zamora and about 9 km to the south, in Reserva Madrigal del Podocarpus, Loja province (Fig 23). We encountered most of the specimens, from Abra de Zamora, outside of Parque Nacional Podocarpus. The species was encountered at an altitudinal range between 2770 and 2960 m a.s.l., in subpa´ramo ecosystems (Fig 2B). Natural history. This is an uncommon species. Lynch [9] described the species based on 10 specimens, and only one more was collected afterwards, in 1984 by W.E. Duellman. Pristiman- tis balionotus is a bromeliad specialist, all the individuals being found inside or on the leaves of terrestrial bromeliads. The call of this species was recorded infrequently in the months of Feb- ruary, May, July, August, September, November and December, only on nights with high humidity (rain or dense fog). All calling males were found on bromeliads leaves. Conservation status. Pristimantis balionotus is currently categorized as Endangered [54] based on criteria B1 ab(iii). We suggest maintaining this category because its Extent of occur- rence (EOO) and Area of occupancy (AOO) are estimated to be less than 30 km2, it is known from only two locations, with part of its populations found outside protected areas which

could be severely affected in the near future by habitat destruction. Remarks. Lynch [9] provides a detailed description of this species. Our diagnosis concurs with all the morphological features described by the author, to which we add descriptions of the characters that are more evident in life. The diagnosis provided herein is based on 10 speci- mens from the original description (KU 142135–44), one adult female (MUTPL 392) and 4 adult males (MUTPL 292, 297, 391, 677) collected from the type locality, and 5 specimens col- lected from Reserva Madrigal del Podocarpus. Pristimantis balionotus is part of the P. phoxocephalus group, but its kinship to the other members of the group remains non-resolved (Fig 21). In our phylogram P. balionotus is closely related to P. hampatusami and an undescribed species (UCS3) but with only moderate BI sup- port. The genetic distance between P. balionotus and its relatives from the P. phoxocephalus group ranges between 2.3–4.4% (P. gloria), 2.5–3.5% (P. lutzae), 3.7–3.9% (P. verrucolatus), 3.6–4.3% (P. tinguichaca), 3.8–5.8% (P. hampatusami) and up to 6.5–8.0% (P. torresi) (S3 Table). Pristimantis versicolor (Lynch, 1979) (Fig 27) Eleutherodactylus versicolor Lynch, 1979 Eleutherodactylus (Eleutherodactylus) versicolor: Lynch, 1996; Lynch and Duellman, 1997 Pristimantis versicolor: Heinicke, Duellman, and Hedges, 2007 Pristimantis (Pristimantis) versicolor: Hedges, Duellman, and Heinicke, 2008 Pristimantis (Huicundomantis) versicolor: Pa´ez and Ron, 2019 Common English name. Variegated Rain Frog (This is a newly proposed name as it is

more appropriate than the previous one, “Loja Robber Frog”). Common Spanish name. Cutı´n versicolor Etymology. The specific name is a Latin adjective meaning of different colors or variegated [25]. Holotype. KU 119858, an adult female [25] obtained 13.5 km E. Loja, 2800 m, a locality just east of the crest on the mountain range dividing Loja and Zamora-Chinchipe provinces, Ecua- dor, on 10 June 1968 by Robert W. Henderson and J. D. Lynch. Paratypes. KU 119859–71 and KU 119911–44, topotypes collected on 10 June and 13 June 1968 by R. W. Henderson and J. D. Lynch. 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 47 / 59