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

Subgenus Huicundomantis Pa´ez and Ron, 2019 Four species of this subgenus were described from Abra de Zamora. Phylogeny For the Huicundomantis subgenus PartitionFinder under AICc (for ML) identified three parti- tion schemes as the best strategy (best model in parentheses): 12S and 16S (GTR+I+G), RAG-1 1st position (HKY+G), and RAG-1 2nd and 3rd positions (TIM+I+G). Under BIC (for BI) Parti- tionFinder identified three partitions: 12S and 16S (GTR+I+G), RAG-1 1st position (K80+G), and RAG-1 2nd and 3rd positions (K81UF+I). The phylogenetic trees constructed by Bayesian inference and Maximum likelihood showed mostly the same topology with differences in the position of some of the unresolved branches as P. atratus, P. totoroi, P jimenezi, and the P. teslai/P. UCS2 subgroup (Fig 21). In general, the branches had stronger support in the case of the Bayesian inference. Similarly to the results of Pa´ez & Ron [30], we recovered the Huicundomantis subgenus as monophyletic with strong support (bootstrap values = 98.7%; posterior probabilities = 1) in both Maximum likelihood and Bayesian analyses (Fig 21). However, in our analysis, the P. cryptomelas group had a strong support only in the case of the Bayesian inference, the ML analysis yielding moderate support. Overall, the phylogenetic tree of our analysis showed the same topology with the one constructed by Pa´ez & Ron [30]. The main differences were in the position of several unresolved branches such as P. jimenezi, P. phoxocephalus, P. totoroi and P.

tinguichaca. Additionally, with the inclusion of the newly generated P. cryptomelas sequences, this species position, relative to P. spinosus, became weakly supported (BI) or only moderately supported (ML) (Fig 21). Our analysis confirmed the membership of P. percultus, P. andinogigas, and P. balionotus to the P. phoxocephalus species group. In the case of P. andinogigas the pairwise genetic distance from its closest congener, P. chomskyi, it was only 2.3% (S3 Table) thus a careful revision of these species statuses is needed. Pristimantis cryptomelas species group One species of this species group was described from Abra de Zamora. Pristimantis cryptomelas (Lynch, 1979) (Fig 22) Eleutherodactylus cryptomelas Lynch, 1979 Eleutherodactylus (Eleutherodactylus) cryptomelas: Lynch, 1996; Lynch and Duellman, 1997 Pristimantis cryptomelas: Heinicke, Duellman, and Hedges, 2007 Pristimantis (Pristimantis) cryptomelas: Hedges, Duellman, and Heinicke, 2008 Pristimantis (Huicundomantis) cryptomelas: Pa´ez and Ron, 2019 Common English name. Cryptic Robber Frog Common Spanish name. Cutı´n crı´ptico Etymology. The specific name is derived from the Greek kryptos, meaning “hidden” and the Greek melas, meaning “black” and refers to the black areas in the axilla and groin and on the hidden surfaces of the limbs [25]. Holotype. KU 141992, an immature female, taken 15 km E Loja, Provincia Zamora- Chinchipe, Ecuador, 2710 m, May 1971 by William E. Duellman. Paratypes. KU 141993, taken syntopically with holotype; KU 120095–96, 8–9 km N San

Lucas, Provincia Loja, Ecuador, 3000 m; USNM 198480–82, Sapote, Provincia Morona-Santi- ago, Ecuador, 2470 m; USNM 198483, 2 km W Sapote, Provincia Morona-Santiago, Ecuador, 2560 m. 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 37 / 59