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Anura, Leptodactylidae) UNIVERSIDADE ESTADUAL PAULISTA “JÚLIO DE MESQUITA FILHO” unesp INSTITUTO DE BIOCIÊNCIAS – RIO CLARO PROGRAMA DE PÓS-GRADUAÇÃO EM CIÊNCIAS BIOLÓGICAS (ZOOLOGIA) Diversidade e evolução do gênero Adenomera Steindachner, 1867 (Anura, Leptodactylidae) CARLA SANTANA CASSINI Tese apresentada ao Instituto de Biociências do Campus de Rio Claro, Universidade Estadual Paulista, como parte dos requisitos para obtenção do título de Doutora em Ciências Biológicas (Área de concentração Zoologia). Março - 2015 CARLA SANTANA CASSINI Diversidade e evolução do gênero Adenomera Steindachner, 1867 (Anura, Leptodactylidae) Orientador: Célio Fernando Baptista Haddad Tese apresentada ao Instituto de Biociências do Câmpus de Rio Claro, Universidade Estadual Paulista, como parte dos requisitos para obtenção do título de Doutora em Ciências Biológicas (Área de concentração Zoologia). Março 2015 CARLA SANTANA CASSINI Diversidade e evolução do gênero Adenomera Steindachner, 1867 (Anura, Leptodactylidae) COMISSÃO EXAMINADORA ____________________________________________________ ____________________________________________________ ____________________________________________________ ____________________________________________________ ____________________________________________________ Rio Claro, 30 de março de 2015. Março 2015 “This is ourselves Under pressure” (Queen, 1981) AGRADECIMENTOS Este trabalho só se concretizou porque ao longo dos anos pude contar com a preciosa ajuda de várias pessoas, às quais agradeço sinceramente: Ao meu orientador Célio pela oportunidade concedida, pelo exemplo profissional e pela confiança depositada em mim. À minha família, Sérvio, Vanira e Pedro pelo estímulo constante e apoio incondicional para que eu seguisse a carreira acadêmica. Aos curadores das coleções José Pombal Júnior, Paulo Garcia, Mirco Solé, Hussam Zaher, Taran Grant e Julián Faivovich pelo empréstimo de tecidos e exemplares sob seus cuidados, sem os quais não este projeto não teria sido viável. Ao Axel Kwet pela atenção nos e-mails e pelos cantos cedidos para análise dos holótipos de A. araucária e A. engelsi, e de exemplares de A. nana e de i sp. n “catarinense”. Ao Phillippe Kok pela gentileza em examinar caracteres morfológicos de Adenomera lutzi e A. heyeri. À Ariadne Fares e Mariana Lyra pela inestimável ajuda no CEIS. Ao Antoine Fouquet pela oportunidade de colaboração. Ao Pedro Taucce pela oportunidade de orientação, colaboração e principalmente por deixar meu dia a dia mais leve. À sua amizade serei sempre grata! À todos que me ajudaram nos trabalhos de campo realizados ao longo desses anos Bianca Berneck e Marina Walker (minha dupla infalível preferida!), Victor Dill, Carlos Paulo Callithrix, Julián Faivovich, William Costa, Iuri Ribeiro, Angele Delieux, Rafel Pontes, Camila Matedi, Carol Calijorne e Thiago Silva Lex. À Mariana Lyra pelo abrigo cedido durante a escrita da tese num ambiente inspirador. Aos colegas de laboratório UNESP-RC Bianca Berneck, Marina Walker, Carol Calijorne, Ariadne Fares, Pedro Taucce, Barnagleison Lisboa, Francisco Brusqueti, Carlos Paulo Callithrix, Fábio Quase Perin, Lucas Bandeira, Danilo Barea Jesus, Eliziane Garcia, Nadya Pupin, Laryssa Teixeira, Vinícius Loredam, Thaís Condez, Délio Baêta, Leo Malagoli, Maria Tereza, Olívia Gabriela e Marília Peri pelo convívio diário produtivo e divertido. Ao Taran Grant e Boris Blotto pelas discussões sempre inspiradoras e produtivas. Ao Victor Dill, por nunca ter se cansado de conversar sobre trabalho comigo, pela compreensão das minhas ausências e por me fazer tão bem mesmo nos momentos mais turbulentos de finais de teses. À sua amizade, companheirismo e amor. MUITO OBRIGADA! Aos meus orientadores de graduação, Renato Feio, e de mestrado, Carlos Cruz e Ulisses Caramaschi, por serem as pessoas que, juntamente com o Célio Haddad, foram os principais responsáveis pela minha formação. Agradeço à Fundação de Amparo à Pesquisa do estado de São Paulo (FAPESP) pelo fomento provido para a realização da tese (através do processo 2010/515606-8) e ao Instituto Brasileiro do Meio Ambiente (IBAMA) e aos diretores dos parques pela licença concedida. Aproveito o momento e agradeço à Comissão de Pós-Graduação pelos prazos concedidos. Sumário INTRODUÇÃO GERAL ........................................................................................................................... 1 REFERÊNCIAS BIBLIOGRÁFICAS ...................................................................................................... 5 CAPÍTULO 1 ......................................................................................................................................... 9 DELIMITAÇÃO ESPECÍFICA DOS CLADOS THOMEI E MARMORATA DO GÊNERO ADENOMERA STEINDACHNER, 1867 (ANURA, LEPTODACTYLIDAE)......................................................................... 9 RESUMO ........................................................................................................................................... 9 ABSTRACT ..................................................................................................................................... 10 INTRODUÇÃO ................................................................................................................................. 11 HISTÓRICO TAXONÔMICO .......................................................................................................... 15 OBJETIVOS ..................................................................................................................................... 20 METODOLOGIA .............................................................................................................................. 21 NATUREZA DOS DADOS E NOMENCLATURA .............................................................................. 21 PROCEDIMENTOS MOLECULARES .............................................................................................. 23 MORFOLOGIA EXTERNA E BIOACÚSTICA .................................................................................. 25 DELIMITAÇÃO ESPECÍFICA ........................................................................................................ 27 RESULTADOS ................................................................................................................................. 28 ANÁLISE FILOGENÉTICA ............................................................................................................ 28 ANÁLISE ACÚSTICA E MORFOLÓGICA ....................................................................................... 38 CARACTERIZAÇÃO TAXÔNOMICA DE ALGUMAS ESPÉCIES DO CLADO MATA ATLÂNTICA DO GÊNERO ADENOMERA ................................................................................................................. 76 DISCUSSÃO .................................................................................................................................. 123 TOPOLOGIA .............................................................................................................................. 123 DELIMITAÇÃO ESPECÍFICA EM ADENOMERA NOS CLADOS THOMEI E MARMORATA ............... 124 ESPÉCIES CRÍPTICAS E ESPÉCIES PARAFILÉTICAS .................................................................... 134 AMOSTRAGEM E UTILIZAÇÃO DE DADOS DIRETOS .................................................................. 136 CONCLUSÕES ............................................................................................................................... 139 REFERÊNCIAS BIBLIOGRÁFICAS .................................................................................................. 140 CAPÍTULO 2 ..................................................................................................................................... 157 EVOLUÇÃO FENOTÍPICA NO GÊNERO ADENOMERA STEINDACHNER, 1867 COM COMENTÁRIOS ACERCA DA SUBFAMÍLIA LEPTODACTYLINAE ................................................................................. 157 RESUMO ....................................................................................................................................... 157 ABSTRACT ................................................................................................................................... 158 INTRODUÇÃO ............................................................................................................................... 159 BREVE HISTÓRICO SOBRE O GÊNERO ADENOMERA .................................................................. 160 FONTES DE EVIDÊNCIA ............................................................................................................ 163 OBJETIVOS ................................................................................................................................... 167 MATERIAIS E MÉTODOS .............................................................................................................. 167 AMOSTRAGEM DOS GRUPOS INTERNO E EXTERNO .................................................................. 167 AMOSTRAGEM DE CARACTERES .............................................................................................. 169 ANÁLISE FILOGENÉTICA .......................................................................................................... 174 RESULTADOS ............................................................................................................................... 176 CARACTERES FENOTÍPICOS ....................................................................................................
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