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Decapoda, Palaemonidae): Uma Abordagem Molecular E Morfológica De Padrões Filogeográficos, Evolução De Características Ecológicas

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Decapoda, Palaemonidae): Uma Abordagem Molecular E Morfológica De Padrões Filogeográficos, Evolução De Características Ecológicas UNIVERSIDADE DE SÃO PAULO FFCLRP - DEPARTAMENTO DE BIOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Sistemática do gênero Palaemon Weber, 1795 (Decapoda, Palaemonidae): uma abordagem molecular e morfológica de padrões filogeográficos, evolução de características ecológicas e status taxonômico das espécies no Brasil FABRÍCIO LOPES DE CARVALHO Tese apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto-USP, como parte das exigências para obtenção do título de Doutor em Ciências - Área: Biologia Comparada. RIBEIRÃO PRETO / SP 2014 UNIVERSIDADE DE SÃO PAULO FFCLRP - DEPARTAMENTO DE BIOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA COMPARADA Sistemática do gênero Palaemon Weber, 1795 (Decapoda, Palaemonidae): uma abordagem molecular e morfológica de padrões filogeográficos, evolução de características ecológicas e status taxonômico das espécies no Brasil FABRÍCIO LOPES DE CARVALHO Tese apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto-USP, como parte das exigências para obtenção do título de Doutor em Ciências - Área: Biologia Comparada. Orientador: Prof. Dr. Fernando Luis Medina Mantelatto Versão Corrigida RIBEIRÃO PRETO / SP 2014 Carvalho, F. L. Sistemática do gênero Palaemon Weber, 1795 (Decapoda, Palaemonidae): uma abordagem molecular e morfológica de padrões filogeográficos, evolução de características ecológicas e status taxonômico das espécies no Brasil 192 p. Tese apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto-USP. Orientador: Fernando Luis Medina Mantelatto 1. Camarão 2. Palaemoninae 3. Filogenia 4. Genética 5. Taxonomia “Nada em biologia faz sentido senão à luz da evolução” Dobzhansky, 1973 “Nada em evolução faz sentido sem uma filogenia” Gould & MacFadden, 2004 Nada faz sentido sem o meu amor, Vanda SUMÁRIO AGRADECIMENTOS..............................................................................................8 PRÓLOGO.................................................................................................................11 RESUMO GERAL...................................................................................................18 GENERAL ABSTRACT........................................................................................19 CAPÍTULO 1. Relações filogenéticas do gênero Palaemon e a evolução de padrões ecológicos e morfológicos................................................................20 RESUMO.....................................................................................................................21 ABSTRACT................................................................................................................22 INTRODUÇÃO.........................................................................................................23 OBJETIVOS...............................................................................................................26 MATERIAL E MÉTODOS..................................................................................27 Obtenção dos espécimes..............................................................................27 Obtenção de dados moleculares...............................................................30 Análises moleculares....................................................................................41 Obtenção de informações sobre hábitat e desenvolvimento larval..................................................................................................................43 Análise morfológica......................................................................................44 RESULTADOS..........................................................................................................47 Relações filogenéticas..................................................................................47 Evolução da ocupação de hábitat e desenvolvimento larval.........50 Morfologia.......................................................................................................51 Palaemon sensu stricto............................................................................52 Linhagem “Alaocaris”.............................................................................53 Palaemon concinnus................................................................................54 5 Palaemon mercedae.................................................................................55 Diferenciação morfológica entre as linhagens de Palaemon sensu lato e a linhagem “Macrobrachium”...................................................................56 DISCUSSÃO...............................................................................................................65 Relações filogenéticas..................................................................................65 Evolução da ocupação de hábitat e desenvolvimento larval.........73 Morfologia.......................................................................................................78 CONCLUSÕES.........................................................................................................86 REFERÊNCIAS........................................................................................................88 CAPÍTULO 2. Uma abordagem molecular e morfológica sobre o status taxonômico das espécies de Palaemon sensu lato no Brasil.....................101 RESUMO...................................................................................................................102 ABSTRACT..............................................................................................................103 INTRODUÇÃO.......................................................................................................104 OBJETIVOS.............................................................................................................109 MATERIAL E MÉTODOS................................................................................110 Obtenção dos espécimes............................................................................110 Análises moleculares..................................................................................110 Análise morfológica....................................................................................122 RESULTADOS........................................................................................................124 Dados moleculares......................................................................................124 Linhagem “Palaemon”..................................................................124 Palaemon mercedae...................................................................124 Palaemon sensu stricto...............................................................124 Linhagem “Alaocaris”...................................................................125 6 Dados morfológicos....................................................................................134 Palaemon argentinus......................................................................134 Palaemon carteri..............................................................................136 Palaemon ivonicus...........................................................................139 Palaemon mercedae.........................................................................141 Palaemon northropi.........................................................................143 Palaemon octaviae...........................................................................147 Palaemon paivai...............................................................................149 Palaemon pandaliformis................................................................151 Palaemon yuna.................................................................................155 Chave para identificação das espécies de Palaemon do Brasil....................................................................................................166 DISCUSSÃO.............................................................................................................169 Linhagem “Palaemon”..............................................................................169 Palaemon mercedae...............................................................................169 Palaemon sensu stricto..........................................................................171 Linhagem “Alaocaris”...............................................................................175 CONCLUSÕES.......................................................................................................182 REFERÊNCIAS......................................................................................................183 7 AGRADECIMENTOS À Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto (FFCLRP) da Universidade de São Paulo (USP), em especial ao Departamento de Biologia e ao Programa de Pós-Graduação em Biologia Comparada, por todos os recursos disponibilizados para a realização de todas as etapas do doutorado. Ao Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) pela bolsa de doutorado (Processo 140199/2011-0) e apoio financeiro (490314/2011-2). À Coordenação Nacional de Pessoal de Nível Superior (CAPES) pela bolsa de doutorado sanduíche no exterior (PDSE Processo 7711-13-1) e verbas do Programa de Apoio à Pós-Graduação (Convênio CAPES/PROAP 56/2007). À Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP Processo 2010/50188-8) pelo apoio financeiro à execução do trabalho. Ao Prof. Dr. Fernando
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