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

vidua, but has a longer call duration, longer inter-call interval, a lower call rate, longer inter- note interval and, most importantly, lower dominant and 90% bandwidth frequency (Table 2). Distribution. Pristimantis samaniegoi is known only from the Cajanuma entrance to the Parque Nacional Podocarpus and about 13 km in the north, in Abra de Zamora (Fig 6). We did not encounter this species in any similar habitats located in the vicinity of the type locality, despite fieldwork being carried out regularly since 2016. The species was encountered at an altitudinal range between 2560 and 3300 m a.s.l., in evergreen upper montane forest (Fig 2C), subpa´ramo (Fig 2B) and shrub pa´ramo ecosystems. Natural history. This is a common species. All the specimens were encountered during the night, close to the ground (usually at 5–20 cm above the ground), at the base of the grassy vege- tation, on the moss bed under the shrubs, or on shrub leaves. Individuals are hard to find, as they are well hidden at the base of the shrubs or inside the moss bed or grassy vegetation. How- ever, the male’s characteristic advertisement call can be commonly heard. Calling males were encountered year round, more frequently on rainy nights. All the females were encountered in the immediate proximity of the calling males. Conservation status. Pristimantis samaniegoi is known only from two close localities, from an estimated area of less than 50 km2. Nonetheless, we recommend that this species be catego- rized as Near Threatened following the IUCN criteria [47], due the fact that it is a relatively

abundant species and its habitat currently does not face any major threats (most of the popula- tion is situated within a nationally protected area—Parque Nacional Podocarpus). Remarks. Pristimantis samaniegoi is part of the P. simonbolivari subgroup, but its kinship to the other members of the subgroup remains non-resolved (Fig 7). In our phylogram P. samaniegoi is most closely related to P. quintanai and an undescribed species, but with only weak BI support. The genetic distance between P. samaniegoi and its closest relatives from the P. simonbolivari subgroup ranges between 4.8–6.0% (P. tiktik) and 6.7–7.3% (P. bambu) (S3 Table). Pristimantis colodactylus (Lynch, 1979) (Fig 15) Eleutherodactylus colodactylus Lynch, 1979 Eleutherodactylus (Eleutherodactylus) colodactylus: Lynch, 1996; Lynch and Duellman, 1997 Pristimantis colodactylus: Heinicke, Duellman, and Hedges, 2007 Pristimantis (Pristimantis) colodactylus: Hedges, Duellman, and Heinicke, 2008 Common English name. Stubby Fingered Rain Frog (This is a newly proposed name as it is more appropriate. The previous one “Piura Robber Frog” makes no sense, as the species is known only from its type locality, Abra de Zamora, Ecuador). Common Spanish name. Cutı´n de dedos cortos Etymology. The specific name is derived from the Greek kolos, meaning “stubby” and the Greek daktylo, meaning “toes”, referring to the short fingers and toes of this species [25]. Holotype. KU 142151, an adult female from 13.5 km E Loja, at the crest (Abra de Zamora)

on the frontier between Loja and Zamora-Chinchipe provinces, Ecuador, 2800 m, obtained on 22 July 1971 by William E. Duellman. Paratypes. KU 142152–59 topotypes; KU 142160–61, 14 km E Loja, Provincia Zamora- Chinchipe, 2770 m; KU 142162–64, 15 km E Loja, Provincia Zamora-Chinchipe, 2710 m. Diagnosis. Pristimantis colodactylus is a small species distinguished by the following combi- nation of traits: (1) skin on dorsum areolate, almost tuberculate (in life the skin areolate texture is more evident); skin on venter areolate; discoidal fold and dorsolateral folds absent; (2) tym- panic membrane and tympanic annulus absent; supratympanic fold absent; (3) snout rounded in dorsal view and in profile; canthus rostralis straight in dorsal view, rounded in profile; (4) upper eyelid bearing one prominent tubercle and several small tubercles, about 62% IOD in 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 28 / 59