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Sistemática E Análise Filogenética De Epiperipatus Clark, 1913 Baseada Em Dados Moleculares E Morfológicos (Onychophora: Peripatidae)

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Sistemática E Análise Filogenética De Epiperipatus Clark, 1913 Baseada Em Dados Moleculares E Morfológicos (Onychophora: Peripatidae) i Capa: Epiperipatus [sp 17] n. sp., por Flávio Uemori Yamamoto Contracapa: papila oral de Eperipatus acacioi, por Cristiano Costa ii Cristiano Sampaio Costa Sistemática e análise filogenética de Epiperipatus Clark, 1913 baseada em dados moleculares e morfológicos (Onychophora: Peripatidae) Systematic and Phylogenetic analysis of Epiperipatus Clark, 1913 (Onychophora: Peripatidae) based on molecular and morphological data Tese apresentada ao Instituto de Biociências da Universidade de São Paulo, para a obtenção de Título de Doutor em Ciências, na Área de Zoologia. Orientador: Dr. Ricardo Pinto da Rocha Coorientador: Dr. Fernando Portella de Luna Marques SÃO PAULO 2016 iii Costa, Cristiano Sampaio Sistemática e análise filogenética de Epiperipatus Clark, 1913 baseada em dados moleculares e morfológicos (Onychophora: Peripatidae) 999 páginas Tese (Doutorado) - Instituto de Biociências da Universidade de São Paulo. Departamento de Zoologia. 1. Onychophora 2. Peripatidae 3. Epiperipatus 4. Revisão sistemática. I. Universidade de São Paulo. Instituto de Biociências. Departamento de Zoologia. Comissão Julgadora: Prof(a). Dr(a). Prof(a). Dr(a). Prof(a). Dr(a). Prof(a). Dr(a) Prof. Dr. Ricardo Pinto da Rocha iv "Without taxonomy to give shape to the bricks, and systematics to tell us how to put them together, the house of biological science is a meaningless jumble" R. M. May, Nature 347:129-130, 1990 v A todos que de alguma maneira contribuíram com meu aprendizado ao longo da vida. vi Agradecimentos Apoio financeiro FAPESP, processos número 2011/20211-0, 2012-02969-6 e 2014/20557-2 Apoio técnico e científico Lab. de Aracnologia, Dep. Zoologia, IB-USP Lab. de Microscopia Eletrônica, MZUSP Lab. de Sistemática Molecular, Dep. Zoologia, IB-USP Giribet Lab, MCZ, Harvard University Lab. de Histologia, Dep. Zoologia, IB-USP Instituições colaboradoras Evolution et Diversité Biologique Laboratory Universidade Federal de Mato Grosso Museu Nacional do Rio de Janeiro/ UFRJ Universidade Federal de Mato Grosso do Sul Museu Nacional da Universidade de São Paulo Universidade Federal de Minas Gerais Museum of Comparative Zoology Universidade Federal da Paraíba Natural History Museum, London Universidade Federal do Piauí Universidad Autonóma de Chiriquí Universidad Nacional de Colombia Participações especiais Aldair de França Neto Orlemir Carrerette dos Santos Fabrício Sanguinetti Cruz de Oliveira Pedro de Souza Dias Manuel Antunes Júnior Renato Mitsuo Nagata Agradecimentos especiais Prof. Dr. Gonzalo Giribet, pelo apoio e ensinamento constante enquanto participei do Giribet lab. Profa. Dra. Cibele Bragagnolo, pelo apoio e sugestões iniciais em meu projeto de doutorado. Prof. Dr. Ricardo Pinto da Rocha, por aceitar o desafio de me orientar e pelas portas abertas. vii Prof. Dr. Fernando Portella de Luna Marques, pela coorientação e ensinamentos de valor incalculável. Prof. Dr. Sérgio Antonio Vanin, pela consultoria jurídica em assuntos nomenclaturais e taxonômicos. Aos alunos do Laboratório de Aracnologia/USP, pelo compartilhamento de conhecimento e amizade. Denis Jacob Machado, pelo apoio nos últimos meses do meu doutorado e por sua dissertação que me serviu de inspiração. Tarciana Barbosa, por todo apoio dedicado e o acompanhamento de cada passo. viii ÍNDICE CAPÍTULO 1. INTRODUÇÃO GERAL .................................................................................................................... 1 I – LINHAS GERAIS SOBRE OS ONICÓFOROS ........................................................................................ 1 II – HISTÓRICO DE EPIPERIPATUS ....................................................................................................... 2 III – DELIMITAÇÃO DE EPIPERIPATUS ................................................................................................. 9 IV – SÍNTESE DOS OBJETIVOS DA TESE ............................................................................................. 11 V – ORGANIZAÇÃO DA TESE ............................................................................................................ 12 VI – REFERÊNCIAS .......................................................................................................................... 13 CHAPTER 2. PHYLOGENETIC STUDIES OF EPIPERIPATUS CLARK, 1913 ............................................................ 17 ABSTRACT ....................................................................................................................................... 17 INTRODUCTION ................................................................................................................................ 18 MATERIAL AND METHODS ................................................................................................................ 19 RESULTS .......................................................................................................................................... 25 DISCUSSION ..................................................................................................................................... 31 SUGGESTED TAXONOMIC ACTS ........................................................................................................ 36 ACKNOWLEDGEMENTS. ................................................................................................................... 40 REFERENCES.................................................................................................................................... 41 SUPPLEMENTARY INFORMATION ...................................................................................................... 46 CHAPTER 3. THE TAXONOMY OF EPIPERIPATUS CLARK, 1913 ........................................................................ 67 ABSTRACT ....................................................................................................................................... 67 INTRODUCTION ................................................................................................................................ 68 A short background. ................................................................................................................................ 68 General introduction ............................................................................................................................... 68 MATERIAL AND METHODS ............................................................................................................... 70 RESULTS AND DISCUSSION ............................................................................................................... 71 Family Peripatidae Evans, 1901 .............................................................................................................. 75 Genus Epiperipatus Clark, 1913 ............................................................................................................. 75 1) Epiperipatus edwardsii (Blanchard, 1847) ......................................................................................... 76 2) Epiperipatus imthurni (Sclater, 1888) ................................................................................................. 81 ix 3) Epiperipatus trinidadensis (Sedgwick, 1888) ..................................................................................... 82 4) Epiperipatus brasiliensis (Bouvier, 1899) .......................................................................................... 83 5) Epiperipatus ohausi (Bouvier, 1900) .................................................................................................. 85 6) Epiperipatus isthmicola (Bouvier, 1902) ............................................................................................ 91 7) Epiperipatus biolleyi (Bouvier, 1902) ................................................................................................. 99 8) Epiperipatus evansi (Bouvier, 1904)................................................................................................. 103 9) Epiperipatus betheli (Cockerell, 1913) ............................................................................................. 104 10) Epiperipatus vespuccii Brues, 1914 ................................................................................................ 106 11) Epiperipatus vagans (Brues, 1925) ................................................................................................. 108 12) Epiperipatus acacioi (Marcus & Marcus, 1955) ............................................................................. 113 13) Epiperipatus barbadensis (Froehlich, 1962) ................................................................................... 121 14) Epiperipatus tucupi (Froehich, 1968) ............................................................................................. 123 15) Epiperipatus machadoi (Oliveira & Wieloch, 2005) ...................................................................... 124 16) Epiperipatus cratensis Brito, Pereira, Ferreira, Vasconscellos & Almeida, 2010 .......................... 133 17) Epiperipatus diadenoproctus Oliveira, Lacorte, Fonseca, Wieloch & Mayer, 2011 ...................... 142 18) Epiperipatus adenocryptus Oliveira, Lacorte, Fonseca, Wieloch & Mayer, 2011 ......................... 144 19) Epiperipatus paurognostus Oliveira, Lacorte, Fonseca, Wieloch & Mayer, 2011 ......................... 146 20) Epiperipatus simoni (Bouvier, 1899) .............................................................................................. 148 21) Epiperipatus
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