ISSN 0066-7870 Arquivos de Zoologia VARIATION AND TAXONOMIC CLARIFICATION OF THE LARGE SPECIES OF THE LEPTODACTYLUS PENTADACTYLUS SPECIES GROUP (AMPHIBIA: LEPTODACTYLIDAE) FROM MIDDLE AMERICA, NORTHERN SOUTH AMERICA, AND AMAZONIA W. Ronald Heyer VOLUME 37, FASCÍCULO 3 MUSEU DE ZOOLOGIA DA UNIVERSIDADE DE SÃO PAULO SÃO PAULO, 2005 UNIVERSIDADE DE SÃO PAULO MUSEU DE ZOOLOGIA Prof. Dr. Adolpho José Melfi Prof. Dr. Carlos Roberto Ferreira Brandão Reitor Diretor Prof. Dr. Hélio Nogueira da Cruz Profª. Drª. Eliana Marques Cancello Vice-Reitor Vice-Diretora Prof. Dr. Adilson Avansi de Abreu Pró-Reitor de Cultura e Extensão Universitária COMISSÃO EDITORIAL Hussam El Dine Zaher (editor) EDITORES ASSOCIADOS Antonio Carlos Marques (Universidade de São Paulo) Carlos José Einicker Lamas (Universidade de São Paulo) Mário de Pinna (Universidade de São Paulo) Sérgio Antonio Vanin (Universidade de São Paulo) SERVIÇO DE BIBLIOTECA E DOCUMENTAÇÃO Teresa Beatriz Nunes Guimarães (bibliotecária) Adriana Nascimento Flamino (bibliotecária) Airton de Almeida Cruz (arte-finalista) MUSEU DE ZOOLOGIA DA UNIVERSIDADE DE SÃO PAULO Av. Nazaré, 481, Ipiranga, CEP 04263-000, São Paulo, SP, Brasil Arquivos de Zoologia. v.15- 1967- Museu de Zoologia da Universidade de São Paulo. v. ilust. ISSN 0066-7870 Continuação de: Arquivos de Zoologia do Estado de São Paulo, v.1-14; 1960-66. 1. Zoologia. I. Museu de Zoologia, USP. As instruções aos autores são publicadas em inglês e português no fascículo introdutório de cada volume, assim como a lista dos consultores que assessoram a Comissão Editorial. Arquivos de Zoologia MUSEU DE ZOOLOGIA DA UNIVERSIDADE DE SÃO PAULO CONTEÚDO W. RONALD HEYER VARIATION AND TAXONOMIC CLARIFICATION OF THE LARGE SPECIES OF THE LEPTODACTYLUS PENTADACTYLUS SPECIES GROUP (AMPHIBIA: LEPTODACTYLIDAE) FROM MIDDLE AMERICA, NORTHERN SOUTH AMERICA, AND AMAZONIA....................................................................................................................................... 269 ISSN 0066-7870 Arq. Zool., S. Paulo São Paulo v. 37 n. 3 p. 269-348 2005 Contents Abstract............................................................................................................................................... 269 Introduction ........................................................................................................................................ 270 Materials and Methods ....................................................................................................................... 270 Discussion ........................................................................................................................................... 337 Acknowledgments .............................................................................................................................. 338 References .......................................................................................................................................... 339 Appendix. Specimens examined. ........................................................................................................ 342 Arquivos de Zoologia MUSEU DE ZOOLOGIA DA UNIVERSIDADE DE SÃO PAULO ISSN 0066-7870 ARQ. ZOOL. S. PAULO 37(3):269-348 30.11.2005 VARIATION AND TAXONOMIC CLARIFICATION OF THE LARGE SPECIES OF THE LEPTODACTYLUS PENTADACTYLUS SPECIES GROUP (AMPHIBIA: LEPTODACTYLIDAE) FROM MIDDLE AMERICA, NORTHERN SOUTH AMERICA, AND AMAZONIA W. R ONALD HEYER1 ABSTRACT Variation of external morphology features and advertisement calls are analyzed for species currently identified as Leptodactylus knudseni, L. labyrinthicus, L. myersi, and L. pentadactylus from Middle America, the Pacific versants of Colombia and Ecuador, northern South America, greater Amazonia, and the corridor of open formations from Argentina to northeastern Brazil. Although there is noticeable geographic variation between eastern and western samples of L. knudseni, the variation is considered to be intraspecific in nature. Geographic variation within L. labyrinthicus is more pronounced and most consistent with recognizing three species: L. labyrinthicus (Spix, 1824), L. turimiquensis new species, and L. vastus A. Lutz, 1930. No new data are available for variation within L. myersi, which had previously been noted as possibly containing two species. Variation within L. pentadactylus is also pronounced and most consistent with recognizing four species: L. pentadactylus (Laurenti, 1768), L. peritoaktites new species, L. rhodomerus new species, and L. savagei new species. Some specimens that had been identified in collections as either L. knudseni or L. labyrinthicus from the Brazilian State of Pará are considered to represent an undescribed species, herein described as L. paraensis new species. Standard simple statistical tests of significance for sexual dimorphism in members of the study group may not indicate biological significance. Adult morphological and advertisement call differentiation patterns among the species recognized in this paper do not provide completely reliable information for identifying the species involved, suggesting a different pattern of differentiation than occurs in the Leptodactylus fuscus species group. Larval morphological variation and habitat differentiation may be important in the evolution of species differentiation in the taxa dealt with in this paper. Key words: Variation, taxonomy, Leptodactylus, Middle America, northern South America, Amazonia. 1 Amphibians and Reptiles, Department of Vertebrate Zoology, National Museum of Natural History, PO Box 37012, Smithsonian Institution, Washington, DC 20013-7012, USA. E-mail: [email protected] Recebido para publicação em 26.01.2004 e aceito em 12.11.2004. 270 Arquivos de Zoologia INTRODUCTION due to problems of storage of large specimens in museum collections. Ulisses Galatti (unpublished manuscript) The limited storage space available did not undertook comparative studies of the reproductive allow me to evaluate variation in the way I have biology of Leptodactylus pentadactylus just north found works most effectively. My optimum method of Manaus in Brazilian Amazonia and central is to borrow all available materials so that as data- Panama. He found striking differences in his studies taking and analysis proceed, specimens can be re- of the organisms at the two sites and convincingly examined when questions arise. For example, concluded that the two populations studied should during data-taking, pattern standards are continually be recognized as distinct species. This conclusion added and specimens evaluated early in the study is supported by the available genetic differentiation often should be re-inspected against the newer data (Maxson and Heyer, 1988). After receiving a standards. draft of Galatti’s unpublished manuscript for The generally small sample sizes of adults comment, I re-examined the data I had taken on available for most localities do not allow a rigorous specimens of L. pentadactylus (Heyer, 1979) to analysis of variation of specimens from single determine whether they were adequate to separate localities, a critical measuring stick for evaluating the specimens I had examined into two taxa; they inter-population variation. were not. Given the nature of the above two problems, At the same time, examination of certain the following approach was used. specimens of large Leptodactylus from the State of Variation in Leptodactylus myersi has been Pará in Brazil indicated that they were neither evaluated relatively recently (Heyer, 1995); L. knudseni nor L. labyrinthicus. I therefore began consequently data for this species are used only to taking more extensive data on large species of compare with data from the other taxa. As noted Leptodactylus in an attempt to define the taxa previously (Heyer, 1995), there is significant involved and to determine their geographic variation within L. myersi that may conform better distributions. The taxa included in this study involve to recognition of two species. There are no new the specimens that were or would be identified as data to address this issue. Leptodactylus knudseni, labyrinthicus, myersi, or Data were taken anew for the following pentadactylus in my previous studies (Heyer, 1979, characters of transformed juveniles and adults: sex; 1995). The large species of the Leptodactylus dorsal pattern; belly pattern; lip pattern; posterior pentadactylus group not included in this study are thigh pattern; upper shank pattern; body folds; male L. fallax, flavopictus, and laticeps, because there arm hypertrophy; male thumb spines; male chest is no evidence to suggest that there are taxonomic spines; male chin, throat, and chest tubercles; upper problems associated with them and they have shank texture; outer tarsal texture; foot texture; distributions largely or entirely apart from the taxa snout-vent length (SVL); head length; head width; included in this study. eye-nostril distance; maximum tympanum diameter including annulus; thigh length; shank length; foot length. MATERIALS AND METHODS Data were accumulated during the years 1994-2003. Evaluating variation to discern species limits Museum abbreviations follow Leviton et al., in the frogs included in this study is made difficult 1985 and Leviton and Gibbs, 1988 (for the two by two problems, both likely related to the large Swedish collections involved), with the exception size of the adults. First, due to space constraints in of OMNH, herein used for the Sam Noble the Division of Amphibians & Reptiles at the Oklahoma