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Análise Fiiogenétíca De Syrphoidea (Ínsecta

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Análise Fiiogenétíca De Syrphoidea (Ínsecta t © © UNIVERSIDADE DO AMAZONAS - UA -«•k INSTITUTO NACIONAL DE PESQUISAS DA AMAZÔNIA - INPA '■•ií o /5ry Ti » Análise Fiiogenétíca de Syrphoidea (ínsecta: 7 1 Diptera) no nível de subfamílias. 7 r\ 7 /•\ 1 r\ r\ 7 o MARIA DENISE DA SILVA MgNEZES } Tese apresentada ao Programa de Pós- r'"-. Graduaçâo em Biologia Tropical e Recursos Naturais do convênio IN PA/U A, como parte dos requisitos para obtenção do título de Doutor em CIÊNCIAS BIOLÓGICAS, área de r' . concentração em ENTOMOLOGIA. r / MANAUS - AM 2001 0 UNIVERSIDADE DO AMAZONAS - ÜA INSTITUTO NACIONAL DE PESQUISAS DA AMAZÔNIA - INPA ■3' 3 •3' BiBLioma 9 3 3 3 "3 O O O Análise Filogenética de Syrphoidea (insecta: ■3 Diptera) no nível de subfamílias. a . ^ <f) •O r^ Maria Denise da Silva Menezes 'TV ; ORIENTADOR: Dr. José Albertino Rafael ''N > ■r\ '/■\ Tese apresentada ao Programa de Pós- Graduação em Biologia Tropical e Recursos Naturais do convênio /-x INPA/UA, como parte dos requisitos para ,f'\ obtenção do título de Doutor em CIÊNCIAS BIOLÓGICAS, área de concentração em ENTOMOLOGIA. MANAUS - AM 2001 ''ÍJ r III ■"> <---- ■ ■ "" ' ~'J ^ \ T-t ■ ■■• ■ ■ ■- • ' -A A /^N ) "'N r\ r \ ^ \ No esforço para entender a diversidade, somos um homem que tenta compreender o mecanismo de um relógio fechado. Ele vê o mostrador e os ponteiros, escuta o tique-tac, mas não tem como abrir a caixa. Sendo habilidoso, poderá imaginar o mecanismo responsável pelo que ele observa, mas nunca poderá estar completamente seguro de que a sua explicação é, realmente, a única possível. Albert Einstein. a\ iv © © o AGRADECIMENTOS %/ © © ,J A Deus. que me tem iluminado os momentos difícieis, amparado nos dias de amargura e conservado a humildade nos momentos de êxito, permitindo-me ír\ ^.r subir mais este degrau, pois sem essa luz, nada conseguiria. <-) Ao Dr. José Albertino Rafael pela confiança e apoio na realização deste O trabalho. ^ A Dra. Rosaly Ale Rocha pelas sugestões e discussões filogenéticas. r\ 3 Ao Dr. Ubirajara Martins, Museu de Zoologia da Universidade de São /-\ . 1 Paulo (MZUSP), pelo empréstimo de parte do material utilizado na análise. O Ao Dr. Jeffrey M. Cumming, Canadian National Collection of Insects (Ottawa. Ontario), pela doação de material utilizado na análise. O Aos Drs. Maria das Graças Vale Barbosa e José Camilo Hurtado O Guerreiro pelo incentivo no início do desenvolvimento do trabalho. O Ao Conselho Nacional de Desenvolvimento Científico e Tecnológico pela r\> concessão da bolsa de estudo. Ao Instituto Nacional de Pesquisas da Amazônia que possibilitou a realização do curso. Aos meus pais e ao Pe. Antonio Cremer, vigário da paróquia de Eirunepé (Am)que ensinaram a perseguir meus objetivos, independente dos dissabores. À Dra. Míriam Silva Rafael, pela amizade. & 9 3 9 A r\ O A r\ r\ 3 n\ . } r\ . y A j 3 r\ y O ) r\ r' . Aos meus filhos, Rafael e Anne Caroline, que são a razão de tudo que faço. Dedico VI o FICHA CATALOGRÁFICA o D MENEZES, Maria Denise da Silva. Análise Filogenética de Syrphoidea (Insecta: Diptera) no nível de subfamílias. Menezes, M. D. S.- Manaus, 2001. V.f r\ 82 p.: il. o Tese de Doutorado - INPA/UA o Syrphoidea - Análise filogenética Pi J J CDD 19^ ed. 595.77 o r\ y '1 O rv rv ' } n Sinopse Análise Filogenética de Syrphoidea (Insecta: Diptera) baseada em caracteres externos e da genitália masculina. Uma hipótese evolutiva é fornecida. Palavras-chave: Syrphoidea (Insecta: Diptera), Análise Filogenética. ^>7 vil SUMARIO 9 9 Agradecimentos Cj Ficha catalográfica vi Sumário vii r\ Lista de Tabelas e Apêndices jx Q r\5 Lista de Figuras Resumo xii r\ Abstract xiii O 7 r\ 1.INTRODUÇÃO 1 7 /f*\ 7 1.1 Considerações sobre a linhagem Syrphoidea + Schizophora 1 r\ . y 1.2 Posição Filogenética de Syrphoidea 1 r\ 1.3' Afinidades internas em Syrphoidea 3 O O -4 Informações gerais sobre Pipunculidae 5 1.4.1 Principais classificações 6 r> 1.5 Informações gerais sobre Syrphidae 7 r\ 1.5.1 Principais classificações 9 t f ] 2. Objetivos 11 ' í / ■ í r 3. MATERIAIS E MÉTODOS 12 1 3.1 Origem do material 12 3.2 Métodos e preparação para estudo 13 3.3 Exame dos espécimes 14 3.4 Terminologia e abreviaturas 14 3.5 Análise Filogenética 15 ® viii © 4. RESULTADOS E DISCUSSÕES 16 í\ 4.1 Classificação 16 rr% 4.1.1. Syrphoidea 17 O 4.1.2. Família Syrphidae 18 4.1.3. As subfamílias de Syrphidae 20 O 4.1.3.1. Subfamília Microdontinae 20 r) 4.2. Caracteres que suportam o monofiletismo do grupo Pipizinae O + Syrphinae + Eristalinae 21 4.3. Subfamília Pipizinae 22 y 4.4. Caracteres que suportam o monofiletismo do grupo Syrphinae r\ y + Eristalinae 23 y 4.4.1 Subfamília Syrphinae 23 r\ 4.4.2 Subfamília Eristalinae 24 y r\ 4.5 Família Pipunculidae 25 . y 4.5.1 Subfamílias de Pipunculidae 27 r\ 4.5.2 Subfamília Nephrocerinae 27 r-\ 4.5.3 Caracteres que sustentam o monofiletismo do grupo Chalarinae"*^.. 30 O O 4.5.4 Subfamília Chalarinae 31 /''A 4.5.5 Caracteres que sustentam o monofiletismo do grupo Protonephrocerinae ^ 33 4.5.6 Subfamília Protonephrocerinae 33 4.5.7 Subfamília Pipunculinae 34 4.6 Classificação proposta para as subfamílias de Syrphoidea 38 4.7 Semelhanças e diferenças entre esta e outras classificações 39 5. CONSIDERAÇÕES FINAIS 40 6. REFERÊNCIAS BIBLIOGRÁFICAS 41 7. Lista de abreviaturas usadas no texto e figuras 53 o IX LISTA DE TABELAS E APÊNDICES G G O Apêndice I. Padrão de cores para estruturas homólogas 70 A r\ O Tabela I. Matriz de caracteres do plano-básico das subfamílías e tribos de Cl Syrphoidea. Números de caracteres correspondentes aos números dos caracteres no texto. O = estado piesiomórfico; 1= estado apomórfico 79 V / O ^ / r\ 1 Apêndice II. Relação dos táxons utilizados para análise dos grupos-externos O r\ 80 O rv f'^. ) ) ) ) ) LISTA DE FIGURAS ) ) ) Figs. 1-4. Lonchopteridae, Syrphidae. Arista 56 ) ) ) Figs. 5 -8. Lonchopteridae, Platypezidae, Opetiidae, Syrphidae. Cabeça, vista • lateral Figs. 9-16. Pipunculidae. Cabeça, vista lateral 58 Figs.17-20. Pipunculidae, Syrphidae, Lonchopteridae, Platypezidae. Cabeça, vista lateral; aparelho bucal 59 Flgs.21-23. Pipunculidae. Aparelho bucal 60 Figs.24-28. Pipunculidae. Aparelho bucal 61 Figs.29-30. Syrphidae. Aparelho bucal 62 Figs.31-36. Syrphidae, Pipunculidae. Asas 63 Figs.37-41. Opetiidae, Lonchopteridae, Platypezidae, Syrphidae. Abdome. Vista ventral, lateral e dorsal 64 Figs.42-46. Pipunculidae, Syrphidae. Abdome vista dorsal, ventral e lateral; Abdôme. Macho, fêmea 65 'N Flgs.47-51. Pipunculidae, Syrphidae. Genitália, fêmea vista lateral 66 Figs.52-58. Pipunculidae. Abdome, macho, vista ventral e lateral 67 Qí Xi Q Figs.59-63. Pipunculidae. Abdome, macho, vista ventral e lateral 68 O Figs.64-68. Pipunculidae, Syrphidae. Abdome, macho, vista ventral, dorsal e lateral 69 O O r\ Figs.69-72. Opetiidae, Platypezidae, Pipunculidae. Genitália, macho 71 > Figs.73-78. Pipunculidae. Genitália, macho 72 » f"\ C) Figs.79-82. Pipunculidae. Genitália, macho 73 O Fígs.83-87. Pipunculidae. Genitália, macho 74 . y Fígs.88-92. Syrphidae. Genitália, macho 75 Figs.93-97. Syrphidae. Genitália, macho 76 S 3 T) Figs.98-100. Syrphidae. Genitália, macho 77 Fig. 101. Cladograma das relações filogenéticas das subfamílias de Syrphoidea. Comprimento: 479 passos; índice de consistência 0,94 78 V,>- o» Xll RESUMO 0 ® Apresenta-se um estudo filogenético de oito subfamílias de Syrphoidea baseado em caracteres externos, aparelho bucal e genítália masculina, inclusive caracteres ainda não utilizados na taxonomia clássica do O grupo. Syrphidae passa a possuir quatro subfamílias, com a seguinte hipótese r\, filogenética: (Microdontinae (Pipizinae (Syrphinae (Eristalinae)))). Os gêneros utilizados na análise foram: Microdon Meigen, Paragodon Thompson, Pipiza Fallén, Pipizella Meigen, Syrphus Fabricius, Toxomerus Macquarf, Paragus r\ Latreille, Baccha Fabricius, Platycheirus Lepeletier & Serville, Melanostoma ■> Schiner, Cheilosa Meigen, Volucella Geoffroy, Ensfa//s Latreille, Eumerus r\ f\ Meigen, Merodon Meigen. Pipunculidae também passa a possuir quatro j r\ subfamílias, com a seguinte hipótese filogenética: (Nephrocerinae (Chalarinae r 7 (Protonephrocerinae (Pipunculinae)))). Os gêneros utilizados na análise foram: 7 Nephrocerus Zetterstedt, Verralia Mik, Chalarus Walker, Jassidophaga Aczél, r\ Protonephrocerus Coilin, Pipuncuius Latreille, Cephalops (Cephalops) Fallén, o Cephalops (Semicephalops) De Meyer, Cephalosphaera (Cephalosphaera) Enderlein, Cephalosphaera (Neocephalosphaera) De Meyer, Dorylomorpha Aczél, Tomosvaryeíia Aczél, Microcephalops De Meyer, Metadorylas Rafael, í - / Elmohardyia Rafael, Allomethus Hardy, Eudoryias Aczél. Todas as subfamílias apresentam caracteres apomórficos que estabelecem o monofiletismo, exceto Eristalinae que aparentemente é um grupo parafilético. A análise foi realizada através da metodologia hennigiana, utilizando o Programa Hennig 86 (Farris, 1988) auxiliado pelo Tree Gardener 2.2 versão Windows, baseada em 58 caracteres. Um cladograma mostrando as hipotéticas relações filogenéticas no nível de subfamílias é fornecido. % xííi ^ ABSTRACT I % % ^ A phylogenetic study of eight Syrphoidea subfamilies based on externai \ characters, mouth parts and male genitalia, including which have not been used on this groups classicai taxonomy yet. According to the foilowing phylogeneticai Syrphidae comes to possess four subfamilies: (Microdontinae ^ (Pipizinae (Syrphinae (Eristalinae)))). The genera used in analysis where: Microdon Meigen, Paragodon Thompson, Pipiza Fallén, Pipizella Meigen, Syrphus Fabricius, Toxomerus Macquarf, Paragus Latreiile, Baccha Fabricius, Platycheirus Lepeietier & Serviile, Melanostoma
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