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Universidade Federal Do Paraná UNIVERSIDADE FEDERAL DO PARANÁ MARCEL KRUCHELSKI TSCHÁ TAXON PULSE: UM MODELO PARA A DIVERSIFICAÇÃO GENÉTICA DE PEIXES EM BACIAS HIDROGRÁFICAS COSTEIRAS CURITIBA 2016 UNIVERSIDADE FEDERAL DO PARANÁ MARCEL KRUCHELSKI TSCHÁ TAXON PULSE: UM MODELO PARA A DIVERSIFICAÇÃO GENÉTICA DE PEIXES EM BACIAS HIDROGRÁFICAS COSTEIRAS Tese apresentada à Coordenação do curso de Pós-Graduação em Ciências Biológicas - Zoologia, da Universidade Federal do Paraná, como requisito parcial para obtenção do título de Doutor em Ciências na área de Zoologia. Orientador: Prof. Dr. Walter A. Boeger CURITIBA 2016 Ao constante sonho de dignidade, justiça e laicidade dedico iii AGRADECIMENTOS Sou lisonjeado pela esmera orientação do Prof. Dr. Walter A. Boeger. O aprimoramento do conhecimento e compreensão de novos conceitos ocorreu graças a suas explanações e sugestões. Agradeço ao convívio de uma década desde o início de minha carreira. Agradeço também ao Dr. Vinícuis Abilhoa pela coorientação e auxílio taxonômico, ao Dr. Daniel R. Brooks pelas explicações e correções acerca do modelo evolutivo de Taxon Pulse e ao Dr. Lutz Bachmann pela recepção na Noruega, paciência e perspicazes sugestões. Agradeço a equipe do Laboratório de Ecologia Molecular e Parasitologia Evolutiva da Universidade Federal do Paraná e ao pessoal do Museu de História Natural da Universidade de Oslo pela vivência e coleguismo. Aos professores e colegas do curso de pós-graduação em Ciências Biológicas - Zoologia, bem como aos funcionários do Departamento de Zoologia da UFPR pelo apoio e aprendizado durante os quatro anos que se completam. À majestosa e inigualável presença da minha querida esposa Tamyres, agradeço pela paciência, compreensão e parceria. Aos meus queridos pais Elza Kruchelski e Paulo Tschá pelo investimento, educação, dedicação e carinho afetuoso. À presença marcante do irmão Mateos K. Tschá. Aos meus valorosos e seletos amigos, biólogos ou não, todos memoráveis. À Adriano Hauer pela companhia e esforço de coleta no litoral, bem como ao povo caiçara das baías de Laranjeiras e Antonina que colaboraram de certa forma. À Universidade Federal do Paraná, ao Departamento de Zoologia e ao Programa de Pós-Graduação pela infraestrutura e oportunidade. E ao CNPq pela concessão da bolsa de estudo, aporte financeiro essencial. iv SUMÁRIO LISTA DE FIGURAS ......................................................................................... vii LISTA DE TABELAS .......................................................................................... x RESUMO............................................................................................................ xi ABSTRACT ....................................................................................................... xii PRÓLOGO ....................................................................................................... xiii INTRODUÇÃO GERAL .................................................................................... 14 Referências ................................................................................................ 18 CAPÍTULO I ..................................................................................................... 21 Sea-level variation influences the demographic history of estuarine and freshwater fishes of the coastal plain of Paraná, Brazil. ................................ 21 Abstract ...................................................................................................... 22 Introduction ................................................................................................ 23 Materials and Methods............................................................................... 25 Results ....................................................................................................... 28 Discussion ................................................................................................. 33 Acknowledgments ...................................................................................... 35 References ................................................................................................ 35 CAPÍTULO II .................................................................................................... 40 Connection and isolation of watersheds affecting the diversification and distribution of genetic variability of coastal freshwater fishes. ....................... 40 Abstract ...................................................................................................... 41 Introduction ................................................................................................... 42 Materials and Methods............................................................................... 44 Results ....................................................................................................... 49 Discussion ................................................................................................. 53 Acknowledgments ...................................................................................... 60 References ................................................................................................ 60 CAPÍTULO III ................................................................................................... 67 Oscillation in phylogeography of coastal freshwater fishes: setting the stage for diversification by Taxon Pulse. ...................................................................... 67 Abstract ...................................................................................................... 68 Introduction ................................................................................................ 69 v Materials and Methods............................................................................... 72 Results ....................................................................................................... 80 Discussion ................................................................................................. 86 Acknowledgments ...................................................................................... 94 References ................................................................................................ 94 Supplementary material ........................................................................... 102 CONSIDERAÇÕES FINAIS ........................................................................... 106 Referências .............................................................................................. 108 vi LISTA DE FIGURAS CAPÍTULO I. Sea-level variation influences the demographic history of estuarine and freshwater fishes of the coastal plain of Paraná, Brazil. Figure 1. Sampling sites in highlands plateaus and coastal plain of Paraná State, Brazil. The numbered circles and arrows represent the sampling sites according Table I. Below, an orthographic projection (left) and a top view (right) illustrate the significant elevation differences between these areas. S.M.: Serra do Mar mountains, C.P.: coastal plain. ................................... 26 Figure 2. The past demographic by the Bayesian skyline plot for species in coastal plain (A and B) and highlands plateaus (C). The box under represents sea-level in the same time scale in the region and the gray bands show interglacial periods (D). Mismatch distributions are shown beside their skyline plot groups.. ............................................................................................... 32 CAPÍTULO II. Connection and isolation of watersheds affecting the diversification and distribution of genetic variability of coastal freshwater fishes. Figure 1. Sampling sites for the three species in the six target watersheds around the bay of Antonina (sites 1, 2, and 3) and bay of Laranjeiras (sites 4, 5, and 6), both minor extensions of the great bay of Paranaguá, Paraná, Brazil. The western basins are depicted in green and the northern basins in blue colors (for details see Table 1). .......................................................... 46 Figure 2. Haplotype networks for Mimagoniates microlepis, Rineloricaria sp., and Scleromystax barbatus in the six target watersheds ........................... 50 Figure 3. Mismatch distribution for Mimagoniates microlepis, Rineloricaria sp., and Scleromystax barbatus. Colors match the watersheds depicted in Figure 1. ..................................................................................................... 52 Figure 4. Demographic histories of populations of Mimagoniates microlepis, Rineloricaria sp. and Scleromystax barbatus reconstructed by Bayesian Skyline Plots. Thick lines represent means and colored areas the 95% vii confidence distribution of the effective population size. Colors match the watersheds depicted in Figure 1. All different graphs have the same x- and y-axis scales. ............................................................................................. 52 Figure 5. Haplotype accumulation curves for Mimagoniates microlepis, Rineloricaria sp. and Scleromystax barbatus. Standard deviation in black lines indicates sampling curve for northern watersheds, and standard deviation in gray lines represents sampling curve for western watersheds.54 Figure 6. Bathymery in the bay of Paranaguá, Brazil. ................................ 59 CAPÍTULO III. Oscillation in phylogeography of coastal freshwater fishes: setting the stage for diversification by Taxon Pulse. Figure 1. Sampling sites for Ancistrus
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