Systematics of the Neotropical Genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylogeny, the Relevance of Non-molecular Evidence, and Species Accounts Author(s): Rafael O. de Sá, Taran Grant, Arley Camargo, W. Ronald Heyer, Maria L. Ponssa, Edward Stanley Source: South American Journal of Herpetology, 9():S1-S100. 2014. Published By: Brazilian Society of Herpetology DOI: http://dx.doi.org/10.2994/SAJH-D-13-00022.1 URL: http://www.bioone.org/doi/full/10.2994/SAJH-D-13-00022.1 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. 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Ponssa5, Edward Stanley1,6 1 Department of Biology, University of Richmond, Richmond, Virginia, 23173, USA. 2 Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, CEP 05508-090, São Paulo, SP, Brazil. Email: [email protected] 3 Current Address: Centro Universitario de Rivera, Universidad de la República, Rivera 40000, Uruguay. Email: [email protected] 4 Department of Vertebrate Zoology, Smithsonian Institution, Washington, DC 20560, USA. Email: [email protected] 5 Fundación Miguel Lillo, San Miguel de Tucumán, Tucumán, Argentina. Email: [email protected] 6 Herpetology, California Academy of Sciences, Herpetology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, California 94118. Email: [email protected] * Corresponding Author. Email: [email protected] Abstract. A phylogeny of the species-rich clade of the Neotropical frog genus Leptodactylus sensu stricto is presented on the basis of a total evi- dence analysis of molecular (mitochondrial and nuclear markers) and non-molecular (adult and larval morphological and behavioral characters) sampled from > 80% of the 75 currently recognized species. Our results support the monophyly of Leptodactylus sensu stricto, with Hydrolaetare placed as its sister group. The reciprocal monophyly of Hydrolaetare and Leptodactylus sensu stricto does not require that we consider Hydrolaetare as either a subgenus or synonym of Leptodactylus sensu lato. We recognize Leptodactylus sensu stricto, Hydrolaetare, Adenomera, and Lithodytes as valid monophyletic genera. Our results generally support the traditionally recognized Leptodactylus species groups, with exceptions involving only a few species that are easily accommodated without proposing new groups or significantly altering contents. The four groups form a pectinate tree, with the Leptodactylus fuscus group diverging first, followed by the L. pentadactylus group, which is sister to the L. latrans and L. melanonotus groups. To evaluate the impact of non-molecular evidence on our results, we compared our total evidence results with results obtained from analy- ses using only molecular data. Although non-molecular evidence comprised only 3.5% of the total evidence matrix, it had a strong impact on our total evidence results. Only one species group was monophyletic in the molecular-only analysis, and support differed in 86% of the 54 Leptodac- tylus clades that are shared by the results of the two analyses. Even though no non-molecular evidence was included for Hydrolaetare, exclusion of that data partition resulted in that genus being nested within Leptodactylus, demonstrating that the inclusion of a small amount of non-molecular evidence for a subset of species can alter not only the placement of those species, but also species that were not scored for those data. The evolu- tion of several natural history and reproductive traits is considered in the light of our phylogenic framework. Invasion of rocky outcrops, larval oophagy, and use of underground reproductive chambers are restricted to species of the Leptodactylus fuscus and L. pentadactylus groups. In con- trast, larval schooling, larval attendance, and more complex parental care are restricted to the L. latrans and L. melanonotus groups. Construction of foam nests is plesiomorphic in Leptodactylus but their placement varies extensively (e.g., underground chambers, surface of waterbodies, natu- ral or excavated basins). Information on species synonymy, etymology, adult and larval morphology, advertisement call, and geographic distribu- tion is summarized in species accounts for the 30 species of the Leptodactylus fuscus group, 17 species of the L. pentadactylus group, eight species of the L. latrans group, and 17 species of the L. melanonotus group, as well as the three species that are currently unassigned to any species group. Keywords. Behavior; Distribution; Life history; Morphology; Taxonomy; Vocalization. Resumen. Se presenta una filogenia del género Leptodactylus, un clado neotropical rico en especies, basada en análises combinados de datos moleculares (marcadores nuclear y mitocondriales) y no moleculares (caracteres de la morfología de adultos y larvas así como de comportamien- to) se muestrearon > 80% de las 75 especies reconocidas. Los resultados apoyan la monofília de Leptodactylus sensu stricto, con Hydrolaetare como su grupo hermano. La monofília recíproca de Hydrolaetare y Leptodactylus no requiere considerar a Hydrolaetare como un subgénero o si- nónimo de Leptodactylus sensu lato. Se reconocen Leptodactylus sensu stricto, Hydrolaetare, Adenomera y Lithodytes como géneros monofiléticos válidos. Los resultados en general resuelven los grupos tradicionalmente reconocidos de Leptodactylus, con excepciones de algunas especies que son reasignadas sin la necesidad de proponer nuevos grupos o alterar significativamente el contenido de los grupos tradicionales. Los cuatro grupos de especies forman una topología pectinada donde el grupo de L. fuscus tiene una posición basal, seguido por el grupo de L. pentadactylus que es el grupo hermano al clado formado por los grupo de L. latrans y L. melanonotus. Se estimó el impacto de los datos no moleculares en los resultados, comparándose los resultados de evidencia total con los de los análises de datos moleculares solamente. Los datos no moleculares representan un 3.5% de la matriz de evidencia total, pero estos datos tuvieron un impacto significativo en los resultados del análisis de eviden- cia total. En el análisis estrictamente molecular solamente un grupo de especies resultó monofilético, y el apoyo difirió en 86% de los 54 clados de Leptodactylus compartidos entre los dos análises. A pesar que datos no moleculares no fueron incluídos para Hydrolaetare, la exclusión de evidencia no molecular resultó en el género estar dentro de Leptodactylus, demostrando que la inclusión de evidencia no molecular pequeña para un subgrupo de especies altera no solamente la posición topológica de esas especies, sino tambien de las especies para las cuales dichos datos no fueron codificados. La evolución de patrones de historia natural y reprodución se evalúan en el contexto filogenético. La invasión de aflora- mientos rocosos y la construción de cámaras de reprodución subterraneas está limitada a los grupos de Leptodactylus fuscus y L. pentadactylus, mientras que la oofagia larval está restringida al grupo de L. pentadactylus. Por otro lado, los cárdumenes larvales, la proteción del cárdumen, y otros comportamientos parentales complejos carecterizan al clado formado por los grupos de especies de L. latrans y L. melanonotus. Los resú- menes de especies incluyen información de sinonímias, etimología, morfología de adultos y larvas, cantos, y distribución geográfica para las 30 especies del grupo de Leptodactylus fuscus, 17 especies del grupo L. pentadactylus, ocho especies del grupo de L. latrans, 17 especies del grupo de L. melanonotus, así como para las tres especies que actualmente no se encuentran asociadas a ninguno de los grupos de especies. Submitted: 07 July 2013 Handling Editor: Marcio Martins Accepted: 18 July 2014 doi: 10.2994/SAJH-D-13-00022.1 South American Journal of Herpetology, 9(Special Issue 1), 2014, S1–S128 CONTENTS INTRODUCTION ........................................................................................................................................................................S4 MATERIALS AND METHODS ...................................................................................................................................................S6 Ingroup delimitation and outgroup sampling .....................................................................................................................S6