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Molecular Phylogenetics and Taxonomy of the Andean Genus Lynchius Hedges, Duellman, and Heinicke 2008 (Anura: Craugastoridae) Author(S): Ana P

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Molecular Phylogenetics and Taxonomy of the Andean Genus Lynchius Hedges, Duellman, and Heinicke 2008 (Anura: Craugastoridae) Author(S): Ana P Molecular Phylogenetics and Taxonomy of the Andean Genus Lynchius Hedges, Duellman, and Heinicke 2008 (Anura: Craugastoridae) Author(s): Ana P. Motta, Juan C. Chaparro, José P. Pombal, Jr., Juan M. Guayasamin, Ignacio De la Riva, and José M. Padial Source: Herpetological Monographs, 30(1):119-142. Published By: The Herpetologists' League DOI: http://dx.doi.org/10.1655/HERPMONOGRAPHS-D-16-00002 URL: http://www.bioone.org/doi/full/10.1655/HERPMONOGRAPHS-D-16-00002 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Herpetological Monographs, 30, 2016, 119–142 Ó 2016 by The Herpetologists’ League, Inc. Molecular Phylogenetics and Taxonomy of the Andean Genus Lynchius Hedges, Duellman, and Heinicke 2008 (Anura: Craugastoridae) 1,2,8 3 2 4,5 6 7 ANA P. MOTTA ,JUAN C. CHAPARRO ,JOSE´ P. POMBAL,JR. ,JUAN M. GUAYASAMIN ,IGNACIO DELARIVA , AND JOSE´ M. PADIAL 1 Departamento de Zoologia, Universidade Estadual Paulista Julio´ de Mesquita Filho, 13506-900, Rio Claro, SP, Brazil 2 Departamento de Vertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, 20940-040 Rio de Janeiro, RJ, Brazil 3 Museo de Historia Natural de la Universidad Nacional de San Antonio Abad del Cusco (MHNC), Plaza de Armas s/n (Paraninfo Universitario), Cusco, Peru 4 Colegio de Ciencias Biologicas´ y Ambientales, Universidad San Francisco de Quito, Diego de Robles y Vıa´ Interocea´nica, Cumbaya´, 17-1200-841, Quito, Ecuador 5 Centro de Investigacion´ de la Biodiversidad y Cambio Clima´tico (BioCamb), Ingenierıa´ en Biodiversidad y Recursos Geneticos,´ Facultad de Ciencias de Medio Ambiente, Universidad Tecnologica´ Indoamerica,´ Calle Machala y Sabanilla, Cotocollao, Quito, Ecuador 6 Departmento de Biodiversidad y Biologia Evolutiva, Museo Nacional de Ciencias Naturales, CSIC; C/Jose´ Gutierrez´ Abascal, 2, 28006, Madrid, Spain 7 Section of Amphibians and Reptiles, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, PA 15213, USA ABSTRACT: We infer species relationships within Lynchius, a frog genus with four species distributed along the paramos and cloud forests of the Andes of northern Peru and southern Ecuador, and assess species diversity in light of comparative analyses of anatomical traits and inferred relationships. Phylogenetic analyses rely on ~7000 base pairs of mtDNA and nuDNA sequences aligned using similarity-alignment and tree- alignment and optimized under maximum likelihood and parsimony criteria. Inferred relationships place Lynchius as the sister group of the widespread genus Oreobates and this clade as the sister group of the high Andean genus Phrynopus. Our analyses corroborate the dissimilar species Lynchius simmonsi as part of this clade and place it as the sister group of the remaining species of Lynchius. Parsimony and maximum likelihood analyses differ in the internal relationships of Lynchius with respect to the placement of L. flavomaculatus, L. nebulanastes, and L. parkeri, but support the existence of two unnamed species. External morphological comparisons provide diagnostic characters for the two new species, which are named and described herein. Lynchius tabaconas is sister to L. flavomaculatus and occurs at ~2800 m in the cloud forests of Santuario Nacional Tabaconas-Namballe, Cajamarca, Peru. Lynchius oblitus occurs in the same area but at a higher elevation (~3300 m) and is sister to a clade formed by L. flavomaculatus and L. tabaconas in parsimony analyses and to L. nebulanastes in maximum likelihood analyses. We provide a new diagnosis for each of the six species and for the genus, as well as some natural history notes. Key words: Brachycephaloidea; Lynchius oblitus sp. nov.; Lynchius tabaconas sp. nov.; Oreobates; Paramo; Phrynopus; Terrarana THE WET and cold grasslands and elfin forests of the high of Lynchius discovered, was found in the cloud forests and Andes have been traditionally considered poor in frog paramos of a locality 15 km east of Loja (Ecuador; Parker diversity. Recent surveys in northern Bolivia and in Peru 1938). This locality is now part of the Podocarpus National have nonetheless unraveled unexpected levels of new species Park, but this species is currently known to occur farther of terraranan frogs in different lineages (e.g., De la Riva north and south within Ecuador. Lynchius nebulanastes 2007; Lehr and Catenazzi 2009; Lehr et al. 2012; Chaparro (Cannatella 1984) and L. parkeri (Lynch 1975) were found et al. 2015). Albeit these high-elevation lineages have been in paramos and cloud forests, even in syntopy, but farther historically considered part of the genus Phrynopus (Lynch south, along one of the westernmost ridges of the Peruvian 1975), molecular data reveal a contrasting scenario where Cordillera de Huancabamba. An unnamed species is also they have independent origins (Heinicke et al. 2007; Hedges known to occur in northern Peru (Lehr 2005; Hedges et al. et al. 2008; Padial et al. 2014). According to inferred 2008; Padial et al. 2014) in the paramos and elfin forests of relationships, species from Bolivia and southern Peru have an area intermediate between the southern distribution of L. been placed in the genera Bryophryne, Noblella, and nebulanastes and L. parkeri and the northern distribution of Psychrophrynella, the radiation in the Andes of central Peru L. flavomaculatus. A fourth, and morphologically dissimilar retained the name Phrynopus, and the genus Lynchius— species, L. simmonsi (Lynch 1974), was recently inferred as target of this study—was erected to include three species the sister group of Lynchius and is now considered part of distributed along the paramos and elfin forests of the this genus (Padial et al. 2012, 2014). In contrast to the other Cordillera Oriental in southern Ecuador and Cordillera de species of Lynchius, L. simmonsi occurs at lower elevations Huancabamba in northern Peru (Hedges et al. 2008). (~1800 m) along the cloud forests of the Cordillera del Along the Cordillera de Huancabamba in northern Peru Condor,´ in Ecuador, on the easternmost ridges of the and the southern parts of the Cordillera Oriental of Ecuador Amazonian versant of the Andes. the western slopes are mostly dry and covered by chaparral- Lynchius remains a poorly known Andean clade and, although its phylogenetic relationships and osteology have like forest, while the summit and Amazonian versant are been studied in some detail (e.g., Lynch 1975; Duellman and moist and covered with dense vegetation. Above ~2000 m Lehr 2009; Padial et al. 2014), several aspects of its elevation on the eastern versant, dense cloud forests and systematics and natural history remain scarcely studied. bush-like elfin forests and paramos gradually replace For example, morphological variation within this clade is montane forests. Lynchius flavomaculatus, the first species poorly understood because species were described indepen- dently and comparisons were made within a now obsolete 8 CORRESPONDENCE: e-mail, [email protected] phylogenetic framework (e.g., species of Lynchius were 119 120 Herpetological Monographs 30, 2016 thoughttobepartofPhrynopus, Ischnocnema, and descriptions and examination of museum specimens. Types Oreobates). After the inference of L. simmonsi as part of of all species of Lynchius were examined. this clade, the patterns of distribution and morphological Specimens collected were photographed alive, euthanized variation within the clade of Lynchius changed considerably. with Lidocaine, fixed in 10% formalin, and preserved in 70% Furthermore, the phylogenetic relationships of Lynchius ethanol. Coordinates were obtained by means of a Garmin remain contentious. Molecular phylogenetic analyses have PSMAP 62sc Handheld Navigator Global Positioning System placed Lynchius as either the sister group of Phrynopus and further assessments for geographical data and elevation (Padial et al. 2009), Oreobates (Hedges et al. 2008; Padial et were performed in Google Earth. All geographic coordinates al. 2014), or Oreobates and Phrynopus (Padial et al. 2012). are based on geodetic datum WGS84. In addition to the undescribed species from northern Museum acronyms are those cited by Frost (2016), except Peru mentioned above, recent fieldwork in this region and in that MHNC refers to Museo de Historia Natural de Cusco, southern Ecuador by some of the authors of this study Universidad Nacional de San Antonio Abad del Cusco, revealed the existence of another unnamed species of Cusco, Peru (MHNCP in Frost 2016). Specimens examined Lynchius and provided additional material for comparison are listed in Appendix I. and more rigorous phylogenetic analyses. Here, we present Molecular Phylogenetics the results of our analyses of
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