Anura, Strabomantidae) from Peru, with Comments on the Osteology of the Genus
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Phyllomedusa 7(1):11-24, 2008 © 2008 Departamento de Ciências Biológicas - ESALQ - USP ISSN 1519-1397 A new species of Phrynopus (Anura, Strabomantidae) from Peru, with comments on the osteology of the genus Linda Trueb1 and Edgar Lehr2 1 Division of Herpetology, Biodiversity Research Center, and Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, Kansas 66045-7561, USA. E-mail: [email protected]. 2 Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstrasse 159, D-01109 Dresden, Germany. E-mail: [email protected]. Abstract A new species of Phrynopus (Anura, Strabomantidae) from Peru, with comments on the osteology of the genus. A new, small species of Phrynopus is described from the eastern slopes of the Cordillera Oriental just north of the Río Huallaga in central Peru. The species resembles P. montium, but is distinguished from it and other congeners by its spatulate snout. The anterior end of the maxillary arcade is broadly rounded in dorsal view, and composed of wide premaxillae that bear extraordinarily large alary processes that are deflected posteriorly at an acute angle. The configuration of the premaxillae accounts for the wide, depressed snout in this species, which in other osteological features resembles P. montium. The osteology of Phrynopus montium is described in some detail in comparison to the new species, with the intention of establishing a baseline for future morphological studies and taxonomic descriptions of strabomantine anurans of this and related genera. Keywords: Anura, Strabomantidae, Strabomantinae, species description, Phrynopus montium, osteology, Peru. Introduction with the remaining South American taxa being allocated to Brachycephalidae, and the West In their revision of Brachycephalidae (sensu Indian and Mesoamerican direct-developing Frost et al. 2006), Hedges et al. (2008a) erected anurans belonging to Eleutherodactylidae and a new taxon, Terrarana, for the 843 species of Craugastoridae, respectively (Hedges et al. New World direct-developing frogs, of which 2008a). Phrynopus is one genus. Phrynopus is one of 16 Strabomantid frogs are arranged in two genera and 490 species placed in the large subfamilies. Phrynopus belongs to the large, South American subfamily Strabomantidae, diverse Strabomantinae, which contains 465 species and 10 genera distributed mainly in northwestern South America, where it is most Received 4 June 2008. Accepted 1 July 2008. diverse in Andean regions of Colombia, Distributed September 2008. Ecuador, and Peru. Strabomantines are distin- Phyllomedusa - 7(1), September 2008 11 Trueb and Lehr guished from members of its smaller sister leptodactylid frogs. Table 1 summarizes group, Holoadeninae (25 species in 6 genera), contributions to our osteological knowledge of by the narrow terminal digits on the fingers and these taxa (sensu lato) by Lynch (1971, 1975, toes, and the usual absence of circumferential 1986) and Cannatella (1984). Of the 18 species grooves in holoadenines. Strabomantines of Phrynopus listed as having been described or usually have expanded terminal digits and figured in previous publications, only three are circumferential grooves. These morphological retained in the genus today—Phrynopus differences are not especially trenchant peruanus, the type species of the genus, and P. characters to distinguish among members of this dagmarae and P. kauneorum, two species speciose, widely distributed, and phenotypically recently described by Lehr et al. (2002a,b). challenging anuran clade; thus, it is hardly Lynch (1975) provided a drawing of part of the surprising that occasionally, molecular data are pectoral girdle of the type species, P. peruanus. required for unequivocal allocation to Lynch (1975) also provided illustrations of the subfamily, especially in the absence of compa- bony skull for one other taxon, Niceforonia rative morphological data. montia, which now is placed in Phrynopus. An unintended consequence of this major Thus, the irony of our osteological knowledge reorganization of direct-developing frogs is that of Phrynopus is that we seemed to have a we must reassess our morphological knowledge relatively firm grasp on it 30 years ago; however, of the group, owing to the rearrangement of today, we have available three renderings of the taxa. Phrynopus is a case in point. Currently, 21 bony skull of one taxon, Phrynopus montium, a species are recognized—auriculatus, ayacucho, drawing of a portion of the pectoral girdle of the barthlenae, bracki, bufoides, dagmarae, type species, P. peruanus, and drawings of the heimorum, horstpauli, juninenesis, kauneorum, dentigerous processes of the vomers of P. kotosh, miroslawae, montium, nicoleae, dagmarae and P. kauneorum and the terminal oblivius, paucari, peruanus, pesantesi, phalanges of P. kauneorum. tautzorum, tribulosus, and thompsoni (Chaparro Many collections contain undescribed et al. 2008, Hedges et al. 2008a). Several species, and as Hedges et al. (2008a) observed, species formerly placed in Phrynopus were “… their analyses and resulting classification transferred to other genera. Among these are: must be regarded as an initial effort waiting to bagrecito (Psychrophrynella), brunneus be expanded and refined.” Herein, we (Hypodactylus), columbianus (Niceforonia supplement their classification with the columbiana), cophites (Bryophryne), elasso- description of a new species of strabomantid discus (Hypodactylus), fallaciosus (Hypo- anuran of the genus Phrynopus from Peru. The dactylus), flavomaculatus (Lynchius), laplacai species was discovered in the collections of (synonym of wettsteini), lucida (Hypodactylus), Louisiana State University Museum of Natural nanus (Niceforonia nana), nebulanastes Science and is distinguished from all other (Lynchius), nigrovittatus (Hypodactylus), Phrynopus by its unusually broad, sloping snout parkeri (Lynchius), peraccai (Hypodactylus), supported by an unusual configuration of the pereger (Oreobates), peruvianus (part = maxilla and premaxilla. Noblella peruviana; part = Psychrophrynella sp. nov.), spectabilis (synonym of Pleurodema Materials and Methods marmorata), simonsii (Pristimantis), and wettsteini (Psychrophrynella). Osteological The taxonomy of strabomantid frogs characters have figured significantly in follows that of Hedges et al. (2008a,b). The taxonomic studies of Phrynopus and allied taxa terminology specific to these anurans is beginning with Lynch’s (1971) seminal work on adopted from Lynch and Duellman (1997) Phyllomedusa - 7(1), September 2008 12 A new species of Phrynopus (Anura, Strabomantidae) from Peru Table 1 - Summary of osteological data published in association with the generic name Phrynopus, with the current taxonomic status of the taxon noted. Taxa denoted with an asterisk are included in the strabomantid subfamily Holoadeninae, whereas all others are included in Strabomantinae (Hedges et al. 2008a). Name used in Current name Psublication Comment previous publication Phrynopus Phrynopus dagmarae Lehr et al. (&2002a: fig. 2) vomer Illustration dagmarae osteological characters Phrynopus Phrynopus Lehr et al. (2002b: figs. 2, 3) vomer and Illustration & kauneorum kauneorum terminal phalange osteological characters Phynopus montium Niceforonia montia Lynch (1971: fig. 104) bony skull in 3 Illustrations & views; Lynch (1975) osteological characters Phrynopus Phrynopus peruanus Lsynch (1975: fig. 13, pectoral girdle Osteological character peruanus *Psychrophrynella Phrynopus bagrecito Lynch (1986: fig. 2A, a, dorsal/ventral Osteological characters bagrecito skull; fig. 3A, pectoral girdle; fig. 3C, hyoid) Hypodactylus Phrynopus brunneus Lsynch (1975) Osteological character brunneus Niceforonia Phrynopus Lsynch (1975) Osteological character columbiana columbianus *Bryophryne Phrynopus cophites Lynch (1975: fig. 2C, terminal Osteological characters cophites phalanges; fig. 12, dorsal/ventral skull) Lynchius Phrynopus Lsynch (1975: fig. 9) skull in dorsal view Osteological character flavomaculatus flavomaculatus *Psychrophrynella Phrynopus laplacai Lsynch (1975) Osteological character wettsteini Hypodactylus Phrynopus lucida Cannatella (1984: fig. 4, dorsal/ventral Osteological characters lucida skull) Naiceforonia nana Phrynopus nan Lsynch (1975) Osteological character Lynchius Phrynopus Cannatella (1984: fig. 4, dorsal/ventral Osteological characters nebulanastes nebulanastes skull) Liynchius parkeri Phrynopus parker Lynch (1975: fig. 2A, terminal Illustrations & phalanges; fig. 10, dorsal/ventral skull) osteological characters Hypodactylus Phrynopus peraccai Lsynch (1975) Osteological character peraccai Orreobates pereger Phrynopus perege Lsynch (1975) Osteological character *Noblella Phrynopus peruvianus Lynch (1975: fig. 2B, terminal Illustrations & peruviana phalanges; fig. 3: pectoral girdle; fig. 11 osteological characters dorsal/ventral skull) Oireobates simonsii Phrynopus simonsi Lsynch (1975) Osteological character *Psychrophrynella Phrynopus wettsteini Lsynch (1975) Osteological character wettsteini Phyllomedusa - 7(1), September 2008 13 Trueb and Lehr except that the term “dentigerous processes of Huaylaspampa, above Acomayo, Departamento vomers” is used instead of “vomerine de Huánuco, Peru, obtained by Ted Parker in odontophores.” We follow the definitions of January 1975. conditions of the tympanum by Lynch and Paratype - LSUMZ 32308, juvenile from S Duellman (1997); the otic region was Hualayspampa, Bosque Cutirragra ca. 2744 m dissected in order