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Instituto Nacional De Pesquisas Da Amazônia Coordenação De Biodiversidade Divisão De Entomologia - Dient INSTITUTO NACIONAL DE PESQUISAS DA AMAZÔNIA COORDENAÇÃO DE BIODIVERSIDADE DIVISÃO DE ENTOMOLOGIA - DIENT SISTEMÁTICA DE Kapala Cameron, 1884 (HYMENOPTERA: CHALCIDOIDEA: EUCHARITIDAE) KARINE SCHOENINGER MANAUS, AMAZONAS ABRIL DE 2018 ii KARINE SCHOENINGER SISTEMÁTICA DE Kapala Cameron, 1884 (HYMENOPTERA: CHALCIDOIDEA: EUCHARITIDAE) Orientador: Dr. Marcio Luiz de Oliveira (INPA) Co-orientadores: Dr. John Heraty (University of California – Riverside, EUA) Dr. Javier Torréns (CRILAR – Anillaco, Argentina) Tese apresentada ao Instituto Nacional de Pesquisas da Amazônia, como parte dos requisitos para obtenção do título de Doutora em Ciências Biológicas, área de concentração em Entomologia. MANAUS, AMAZONAS ABRIL DE 2018 iii Tese aprovada como requisito para a obtenção do título de Doutora em Ciências Biológicas, área de concentração em Entomologia, no Programa de Pós-Graduação em Entomologia do Instituto Nacional de Pesquisas da Amazonia, pela comissão formada pelos doutores (as): Drª. Rosaly Ale Rocha Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular Dr. Daniel Rodrigo R. Fernandes Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular Dr. Danilo Pacheco Cordeiro Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular Dr. Edinaldo Nelson dos S. Silva Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular Dr. Ronildo Baiatone Alencar Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular iv S365 Schoeninger, Karine Sistemática de Kapala Cameron, 1884 (Hymenoptera: Chalcidoidea: Eucharitidae) / Karine Schoeninger. - Manaus: [sem editor], 2018. xxi, 280f.: il. color Tese (Doutorado) - INPA, Manaus, 2018. Orientador: Marcio L. Oliveira. Coorientadores: John Heraty, Javier Torréns Área de concentração: Entomologia. 1. Eucharitinae. 2. Complexo furcata. 3. Parasitoides de formigas.. I. Título CDD 595.79 Sinopse: Foi realizada a revisão taxonômica, bem como um estudo filogenético com base em dados morfológicos e moleculares para o gênero Kapala , com ênfase nos complexos de espécies iridicolor e furcata. Foram descritas 15 espécies novas , realizadas 4 sinonímias, designados três léctótipos e realizadas sete associações sexuais. Dois novos gêneros são propostos, Ecariatana n. gen. e Torquata n. gen. Palavras-chave: Eucharitinae, complexo furcata, complexo iridicolor, Região Neotropical, parasitoides de formigas v AGRADECIMENTOS Ao Instituto Nacional de Pesquisas da Amazônia (INPA) e ao Curso de Pós-Graduação em Entomologia, pela infraestrutura e oportunidade de realização do doutorado deste projeto. À Coordenação de Aperfeiçoamento de Pessoal de Nível Superior pelo primeiro ano de concessão de bolsa de doutorado sanduíche (processo PDSE - 88881.133943/2016-01). Ao Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), pelos três últimos anos de concessão de bolsa. Ao meu orientador Marcio L. Oliveira pela orientação e suporte ao longo destes últimos anos. Gostaria de agradecer imensamente aos Drs. John Heraty e Javier Torréns pela co- orietação e apoio ao longo destes quatro anos de doutorado. O zelo de ambos pelo mundo natural e o seu amor por Eucharitidae são aparentes em todos os seus trabalhos. Vocês me encorajaram a pensar de maneira mais criativa e a estabelecer metas elevadas. Eu aprendi muito nos últimos anos com vocês. A cada curador das 52 coleções que enviaram material de estudo e que proporcionaram a realização deste trabalho, em especial a curadora do The Natural History Museum, Dra. Natalie Dale-Skey, por toda a sua ajuda e compreensão. Aos amigos do INPA e colegas do Laboratório de Hymenoptera: Pedro, Breno, Alexandre, Diego, David, Sian, Bruno, Itanna, Thiago, Jeane, Patrik, Nikolas, Alexssandro, Ísis, Daniara, Thaís, Daniel, pelo companheirismo, discussões eternas sobre trabalho e, principalmente filogenia, e pelos momentos de descontração. Desde já, aos membros da banca pelas contribuições e sugestões. Um agradecimento especial aos meus pais, Artélio e Marilene, pelo amor contante e pelo apoio, tanto nos momentos felizes quanto nas dificuldades. Por terem compreendido a minha ausência durante todos esses anos (seis anos até o momento e sei que não foi fácil). Sem vocês eu não teria chegado até aqui. Amo vocês! Minha gratidão sincera a todo o pessoal que me incentivou nesta tarefa. E a todos os que, indiretamente colaboraram para que eu pudesse terminá-la. Obrigada. vi RESUMO Kapala Cameron, 1884 é amplamente difundido em toda a região Neotropical, com exceção do Chile e uma espécie disjunta que é encontrada na África Central e Madagascar. A necessidade de uma revisão das espécies de Kapala foi observada a partir de suas descrições originais, que são extremamente sucintas, o que resultou em muitas identificações errôneas. A grande maioria das descrições menciona a coloração, que é extremamente variável; a presença de dois longos espinhos, presença de antenas flabeladas e mesoscuto alto e convexo. Essas características são comuns a todas as espécies tornando a identificação de suas espécies quase impossível. Apesar disso, a maior dificuldade encontrada dentro do grupo é a sua alta variação instraespecífica, mas ao mesmo tempo altamente conservada. Esta é a primeira tentativa de revisar as espécies conhecidas, descrever novos táxons e estabelecer uma linha de base para a taxonomia do gênero. Além disso, uma nova proposta filogenética para Kapala é fornecida. No Capítulo I, com base no exame dos espécimes-tipo e material complementar, redescrevemos as espécies-tipo do gênero e estabelecemos os limites das espécies válidas; quatro sinônimos foram realizados, K. atrata (Walker, 1862) sin. nov. e K. surgens (Walker, 1862) sin. nov. foram sinonimizadas com K. flabellata (Fabricius, 1804), K. striaticeps (Cameron, 1913) sin. nov. foi sinonimizada com K. inexagens (Walker, 1862) e K. sulcifacies (Cameron, 1904) sin. nov. foi sinonimizado com K. romandii (Guérin-Meneville, 1845); três lectótipos foram designados: K. cynipsea (Walker, 1862) pres. desig., K. inexagens (Walker, 1862) pres. desig. e K. terminalis (Ashmead, 1892) pres. desig., e sete espécies tiveram machos associados a fêmeas. Sete novas espécies são descritas: K. confusa n. sp., K. corcovata n. sp., K. genistriata n. sp., K. gracilispina n. sp., K. haplospinosa n. sp., K. jalisca n. sp. e K. spinaepplanata, e uma chave de identificação para todas as espécies é fornecida. No capítulo II, um estudo filogenético com base em dados morfológicos e moleculares foi realizado com o objetivo de testar a monofilia de Kapala e as relações entre as suas espécies. Para este propósito, 61 caracteres relacionados à morfologia externa foram analisados. Para as análises moleculares, partes de cinco regiões de genes foram amplificadas: 18S, 28S-D2, 28S-D3-D5, COI-nj e COII. Como resultado, foi verificado em todas as análises, morfológica e molecular, que Kapala é parafilético, dividido em três complexos de espécies, que por sua vez, estão divididos por agrupamentos de gêneros distintos. No capítulo III, foi realizada a revisão do complexo de espécies iridicolor, bem como uma filogenia molecular vii com base em 5 genes, 3 nucleares e dois ribossomais. Como resultados, foram reconhecidas seis espécies dentro do complexo iridicolor, incluindo K. iridicolor descrita por Cameron (1904) e cinco novas espécies, sendo elas: K. cavicornis n. sp., K. ceciliae n. sp., K. ipa n. sp., K. longicornis n. sp. e K. parairidicolor n. sp. Além disso, uma diagnose do complexo K. iridicolor é fornecida, bem como uma chave de identificação para as espécies. No capítulo IV, foi realizada a revisão do complexo de espécies furcata e, também, uma filogenia molecular, sendo reconhecidas três novas espécies: K. deltalis n. sp., K. parafurcata n. sp. e K. quasimodo n. sp. Além disso, uma diagnose para o complexo furcata é fornecida, bem como uma chave de identificação para as suas espécies. No capítulo V, como base na filogenia morfológica e molecular, dois novos gêneros são descritos para a família Eucharitidae, Ecarinata n. gen. e Torquata n. gen., sendo fornecida uma chave revisada e atualizada para os gêneros Neotropicais de Eucharitidae. Palavras-chave. Eucharitinae, complexo furcata, complexo iridicolor, neotropical, parasitoides de formigas. viii ABSTRACT Kapala Cameron, 1884 is widespread throughout the Neotropical region, except for Chile and a disjunct species is found in Central Africa and Madagascar. The need for a review of the Kapala species was observed from their original descriptions, which are extremely succinct, resulting in many misidentifications. The vast majority of descriptions mention coloration, which is extremely variable; the presence of two long spines, flabellate antennas in males and mesoscuto high and convex. These characteristics are common to all species making identification of their species almost impossible. Despite this, the greatest difficulty found within the group is its high intraspecific variation, but at the same time highly conserved. This is the first attempt to revise known species, to describe new taxa, and to establish a baseline for genus taxonomy. In addition, a new phylogenetic proposal for Kapala is provided. In Chapter I, based on the examination of the species-type and complementary material, we redescribe the type species of the genus and establish the limits of valid species; four synonyms were performed, K. atrata (Walker, 1862) n. syn. and K. surgens (Walker, 1862) n. syn. was synonimized with K. flabellata (Fabricius, 1804), K. striaticeps (Cameron, 1913) n. syn. was synonymized
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