**Taxonomy**

*Qosqophryne new genus*

http://zoobank.org/urn:lsid:zoobank.org:act:7DDB98AD-CCF9-4977-B814-285D25B3D1BF **Type species.** *Bryophryne gymnotis* Lehr and Catenazzi, 2009

**Included species.** *Qosqophryne flammiventris* (Lehr and Catenazzi, 2010), comb. nov.; *Q. mancoinca* (Mamani, Catenazzi, Ttito, Mallqui, Chaparro, 2017), comb. nov.

**Diagnosis.** (1) Head wider than long, narrower than body, body robust, extremities short; (2) tympanic membrane and annulus present; (3) cranial crests absent; (4) prevomerine teeth and dentigerous process of vomers present (but absent in *Q. flammiventris*); (5) trips of digits narrow, rounded, circumferential grooves absent, terminal phalanges T-shaped to knobbed; (6) Finger I shorter than Finger II, nuptial pads absent; (7) Toe V shorter than Toe III; (8) fingers and toes with lateral fringes (but absent in *Q. flammiventris*); (9) subarticular tubercles small, rounded; (10) dorsolateral folds short, discontinuous or continuous; (11) discoidal fold absent (present in *Q. mancoinca*); (12) trigeminal nerve passing external to *m. adductor mandibulae externus* ('S' condition; Lynch, 1986); (13) snout-vent length from 16.7–19.3 mm in males and 16.0–22.2 mm in females of *Q. gymnotis*, to 19.6–22.9 mm in males and 23.6–26.5 mm in females of *Q. mancoinca*; (14) males with median subgular vocal sac and vocal slits, nuptial pads absent; (15) advertisement call whistle-like, composed of a single, tonal note in *Q. gymnotis*, 2–3 short notes in *Q. mancoinca*, and 3–4 short notes in *Q. flammiventris*.

There are no known morphological synapomorphies for *Qosqophryne*, but the three known species share the following traits (Table 3): (1) males with median subgular vocal sac produce whistle-like tonal calls composed of 1–4 short notes; (2) tongue ovate; (3) skin on venter smooth to weakly areolate (in *Q. flammiventris*); (4) inner tarsal fold absent. Four other genera of Holadeninae occur south of the Apurimac canyon, a proposed biogeographic barrier for high-elevation terrestrial breeding frogs [13–15]. *Bryophryne* di ffers from *Qosqophryne* in lacking an externally visible tympanum, and having males without vocal sac and not emitting vocalizations [2,12,16]. *Oreobates* have head about the same width as body, smooth venter, subarticular and supernumerary tubercles large, conical or subconical, projecting, and range in snout-vent length from 20–63 mm [1,5]. *Noblella* and *Psychrophrynella* have smooth venter, elongated tongue, two prominent metatarsal tubercles, and in most species facial masks and/or a tarsal fold-like, sigmoid tubercle [2,19,20,28]. *Qosqophryne* is most similar to its sister genus *Microkayla*. Putative synapomorphies of *Microkayla* are a rounded tongue, areolate belly, and absence of prominent metatarsal tubercles [2]. It is presumed that all species of *Microkayla* vocalize, and known calls consist of a simple, short whistle-like tonal note [2,4]. *Qosqophryne* di ffers from

most *Microkayla* in having (except for *Q. flammiventris*) fingers and toes with lateral fringes (absent in *Microkayla*), and having (except *Q. flammiventris*) dentigerous processes of vomers (absent in *Microkayla*). Future examination of osteological characters, for example through computed tomography, might help identify such characters, and resolve the condition of the tympanic apparatus in the three genera *Bryophryne*, *Microkayla* and *Qosqophryne*.

**Table 3.** Meristic traits (+ = character present, - = character absent) for the three known species of *Qosqophryne* gen. n.


**Etymology.** The name refers to the city of Cusco, using the spelling Qosqo which more closely reflects the name in Quechua. *Qosqo* is used in apposition with *phryne*, from the greek for "frog". Thus, the name for the new genus alludes to the geographic distribution of the three known species in the Peruvian Department of Cusco.

**Distribution, natural history, and conservation.** The three species of *Qosqophryne* occur within a region of ~150 km<sup>2</sup> in the upper montane forests and grasslands of the Cordilleras de Urubamba and Cordillera de Vilcabamba, Provincia La Convención, Department Cusco, Peru. These frogs inhabit cloud forests, elfin forests, montane scrub and humid grasslands (puna) from 3270 to 3800 m a.s.l. Similar to other regions in the high Andes, these habitats and their amphibian communities are threatened by pasture burning, climate change and associated expansion of agricultural activities, deforestation, and the fungal disease chytridiomycosis [43,44]. Although chytridiomycosis has caused the collapse of montane frog communities at several sites in Departamento Cusco [45,46], terrestrial-breeding frogs have generally declined the least, and several species challenged in experimental infection trials appears to resist or tolerate infection [47]. Protection of natural habitats will benefit conservation of these frogs. Two of the three species occur within naturally protected areas: *Q. gymnotis* within the Área de Conservación Privada Abra Málaga, and *Q. mancoinca* within Machu Picchu Historic Sanctuary.

**Remarks.** The new genus is distinguished from all species of *Bryophryne* by the presence of tympanum and tympanic annulus, and median subgular vocal sacs in males. Furthermore, males of all three species of *Qosqophryne* are known to emit advertisement calls (unknown in all species of *Bryophryne*, except possibly for *B. bustamantei*). We have described the advertisement calls of *Q. gymnotis* and *Q. mancoinca* [14,17]. One of us (LM) has recorded the advertisement call of a male *Q. flammiventris*(MUBI 13365) at the type locality, and this call is composed of 3–4 short notes (~15–35 ms duration) at dominant frequency ~3000 Hz. Females of *Q. gymnotis* attend clutches of 14–16 eggs [39], but unattended clutches of up to 19 eggs have also been found [14].

The new genus *Qosqophryne* is supported by our molecular phylogeny, the most complete to date covering three mitochondrial and two nuclear gene fragments, as well as most described species of *Bryophryne* and *Microkayla*. Despite the absence of known synapomorphies for the sister clades *Microkayla* and *Qosqophryne*, we are confident that our proposed arrangemen<sup>t</sup> reflects the evolutionary history of these organisms, and ye<sup>t</sup> still takes into consideration taxonomic stability [48]. There is strong support (bootstrap probabilities) at the node where *Microkayla* and *Qosqophryne* diverge, and the relative branch lengths leading to their respective living species is similar, or in some cases exceed the branch lengths separating other genera within Terrarana (e.g., *Euparkerella* and *Holoaden*, or *Barycholos* and the "northern clade" of *Noblella*).
