**1. Introduction**

Terrestrial-breeding frogs of the high Andes display an impressive degree of evolutionary convergence [1–4]. Such convergence is associated with life in the cloud forest and high-Andean grassland. Frogs in many genera of Terrarana have evolved strikingly similar body forms [4,5], typically a small, compact body with very short legs and feet, short arms and hands, loss of toe pads and discs, head wider than long, small eyes directed anterolaterally, and, in many groups, reduction or loss of tympanic structure and function [3]. The high similarity of body forms has delayed obtaining a

taxonomic arrangemen<sup>t</sup> that reflects the evolutionary history and phylogenetic relationships of most species of small, terrestrial-breeding frogs of the Andes [1,6,7].

Illustrating the complexity within Terrarana of identifying monophyletic groups in presence of ecological convergence, authors originally assigned frogs belonging to di fferent evolutionary lineages to the genus *Phrynopus* [1,8,9]. Indeed, *Phrynopus* might still contain incorrectly classified species of *Pristimantis* that lack vocal sacs, external tympanic apparatus and toe pads [10]. Subsequent molecular analyses revealed a much greater diversity and deeper genetic structure, such that Hedges et al. [1] proposed to split *Phrynopus* into four genera, and to erect the new subfamily Holoadeninae to include the newly described genera *Bryophryne*, *Niceforonia*, and *Psychrophrynella*. Within Holoadeninae, the molecular phylogeny by Hedges et al. [1] recognized *Bryophryne* as a distinct clade on the basis of DNA sequences from a single species, *B. cophites* (formerly *Phrynopus cophites* Lynch, 1975). Hedges et al. [1] used morphological characters to assign to *Bryophryne* a second species, *Phrynopus bustamantei* Chaparro, De la Riva, Padial, Ochoa, and Lehr, 2007. The new genus *Byrophryne*, along with the other genera of Holoadeninae, was recognized using molecular data, despite the lack of morphological synapomorphies [1,2,5,11].

Since Hedges et al. [1] published their molecular phylogeny, researchers have continued discovering terrestrial-breeding frogs: the number of species of *Bryophryne* has increased from two to 14 species [12–17], and the number of species across all Holoadeninae genera from 36 to 151 species [8]. As far as we know, all species of *Bryophryne* have micro-endemic distribution, and are only known to occur at their respective type localities and immediate surroundings [2,12,14–16,18]. The most recent phylogeny included six of the 14 species of *Bryophryne*, and recovered *Bryophryne* as being the sister taxon to the clade containing *Barycholos*, "*Eleutherodactylus bilineatus*", *Euparkerella*, *Holoaden*, and *Noblella* [2]. However, this phylogeny by De la Riva et al. [2] did not include sequences of the three species of *Bryophryne* having an external tympanum and males with subgular vocal sacs, because sequences were unavailable at the time. Additionally, De la Riva et al. [2] erected a new genus, *Microkayla*, to accommodate all species of *Psychrophrynella* from Bolivia (and one species of *Psychrophrynella* from Peru), as well as two new species from Peru. Because of these discoveries, the integration of molecular, acoustic and morphological approaches, and the ongoing revision of existing and new material, we have a better understanding of the diversity in this group of cryptic genera. As part of our ongoing work, we have become aware of (1) uncertainty regarding the evolutionary relationships of *Noblella* and *Psychrophrynella* [2,19,20], (2) an underestimated species richness and endemism in *Noblella* and *Psychrophrynella* [19–22], and (3) three species of *Bryophryne* (*B. flammiventris*, *B. gymnotis*, *B. mancoinca*; Figure 1) having traits not shared with any other species of *Bryophryne*, such as having an external tympanum and males with subgular vocal sacs and emitting advertisement calls. Here we address the latter of these findings, and propose a new genus for the only three species of *Bryophryne* known to produce vocalizations and possessing external tympanic membrane and annulus.

**Figure 1.** Holotypes of species of *Qosqophryne* gen. n. in dorsolateral and ventral views: (**A**,**B**) *Q. flammiventris* (MUSM 27613; SVL 19.8 mm): (**C**,**D**) *Q. gymnotis* (MUSM 25543; SVL 18.4 mm); (**E**,**F**) *Q. mancoinca* (MUBI 11152; SVL 26.5 mm). Photographs by E. Lehr (**A**,**B**), A. Catenazzi (**C**,**D**) and L. Mamani (**E**,**F**).

## **2. Materials and Methods**

We are familiar with most described species of *Bryophryne*, which we have seen in the field or inspected in collections. We provide a complete list of examined specimens in Appendix A. We used the literature (i.e., original species descriptions) for species whose specimens we could not examine. We have described the advertisement calls of *B. gymnotis* and *B. mancoinca* [14,17], and have heard and provided a short description of the call of *B. flammiventris* [15]. We refer readers to the original publications for details on recording methods.

We combined DNA sequences available from GenBank with sequences from newly collected tissues to generate molecular phylogenies of *Bryophryne* and closely related Holoadeninae taxa (Table 1). We considered sequences for a fragment of the 16S rRNA gene (16S), a fragment of the 12S rRNA gene (12S), the protein-coding gene cytochrome c oxidase subunit I (COI), the nuclear protein-coding gene recombination-activating protein 1 (RAG1), and the tyrosinase precursor (Tyr). All taxa selected for our comparisons belong to the subfamily Holoadeninae [1,23,24].


**Table 1.** GenBank accession numbers for taxa and genes sampled in this study. Genbank accession codes of the new sequences are highlighted in bold font.

**Table 1.** *Cont*.



**Table 1.** *Cont*.
