Sistemática de Bethylinae (Hymenoptera, Bethylidae)
Autor(a) principal: | |
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Data de Publicação: | 2017 |
Tipo de documento: | Tese |
Idioma: | por |
Título da fonte: | Repositório Institucional da Universidade Federal do Espírito Santo (riUfes) |
Texto Completo: | http://repositorio.ufes.br/handle/10/9917 |
Resumo: | The genera of Bethylinae are clearly different one from another. The most of Bethylinae species are known only by the female sex. Their taxonomic boundaries can be considered well established, except for Goniozus Förster. Undoubtedly it is the genus with the highest degree of taxonomic confusion regarding its boundaries. The important characters delimiting Goniozus are also shared by several Bethylinae genera, making classification uncertain and hampering understanding of character evolution and variation between taxa. This subfamily currently comprises approximately 540 species described worldwide classified in eight extant genera with low sexual dimorphism. The phylogeny of Bethylinae lineages has received attention by Sorg in 1988, Polaszek & Krombein in 1994, Terayama in 1995 and De Ploëg & Nel in 2004. In all previous analyses performed there is a basal polytomy among Eupsenella, Lytopsenella and the remaining Bethylinae genera. Here we aimed 1) to review the fossil species of Bethylinae, providing descriptions and illustrations when necessary. In addition, o provide a new key to the fossil species of Bethylinae; 2) to recognize, define and describe Afrobethylus as a new Bethylinae genus, its new species. In addition, to define their diagnostic characteristics, as well as provide a key to species of Afrotropical region based on females; 3) to propose a phylogenetic hypothesis based on morphological data of the genera of Bethylinae drawing especial attention to Eupsenella and Lytopsenella relationships with the other genera of this subfamily; 4) to investigate and discuss the main diagnostic characters of Bethylinae genera. Examined material was provided by several institutions. The descriptions, character list, character matrices for cladistic analysis and key, when necessary, were elaborated with o software DELTA. The searches for the most parsimonious trees were carried out under the software TNT. Here, based on our results, we considered that the fossil genus Protobethylus De Ploëg & Nel, 2004 is a junior synonymous of Eupsenella Westwood, 1874. In addition, based on our rescue and compilation of whole the existing information in Bethylinae the fossil family Fushunochrysidae proposed by Hong in 2002 was synonymized with Bethylidae Haliday, 1839. The single genus of this family is Fushunochrysites Hong, 2002 and its single species F. eocenicus Hong, 2002 was established as its type-species. Here, we propose that the best placement of this monotypic genus in Bethylidae is into Bethylinae. Fushunochrysites displays on its forewing several characters that are also present in all members of Eupsenella. Moreover, Sinibethylus Hong, 2002 from Chinese Xilutian coal mine is here also synonymized with Eupsenella. Finally, the last synonymy of genus proposed here is related to Messoria that was described originally by Meunier in 1916. We here propose 10 Messoria as a new junior synonym of Goniozus and transfer its single Messoria copalina Meunier, 1916 to Goniozus. In addition, based on an unusual combination of characters never seen before to Bethylinae we describe Afrobethylus as a new Bethylinae genus. This Bethylinae genus is the unique excluviley Afrotropical. In the phylogenetic context, the resulting matrices contain a total of 61 species of Bethylinae terminals as ingroup, with a total of 43 characters were coded for males and a total of 87 species, 44 characters were coded for females. The resampling analyses based on both males and females returned support for a sister-group relationship among all extant Bethylinae genera. All genera were retrieved as lineages in both analyses, except Goniozus that was retrieved as paraphyletic in all analyses performed. Our most remarkable result obtained was the resolution of the existing basal polytomy in Bethylinae between Lytopsenella and Eupsenella with the other extant Bethylinae genera. In all analyses performed, these genera were retrieved by the first time as sister-group each other and separated from the other genera in all analyses performed. Based on our results, we discuss the main diagnostic characters of Bethylinae genera mainly present in the antenna, forewing, and finally by the first time in the male and female genitalia. In special, about the forewings in Bethylinae, we suggest practicable approaches to functional morphology to the venation in the Bethylinae clades. The present study is the most comprehensive cladistic treatment dedicated to the understanding of the evolution of the Bethylinae genera, and the first to consider a global sampling of species of this subfamily. We conclude that the morphological characters unexplored, mainly, of male and female genitalia might offer additional data relevant to more robustly estimate the phylogenetic history of this group. Here, we also promote a first step to direct and encourage future research in Bethylinae using the arrangement of forewing veins, flexion lines and male and female genitalia characters to phylogenetic reconstruction. |
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Azevedo, Celso OliveiraRamos, Magno SupraniAguiar, Alexandre PiresMonteiro, Cecília WaichertGobbi, Fernanda ToniniHermes, Marcel Gustavo2017-02-01The genera of Bethylinae are clearly different one from another. The most of Bethylinae species are known only by the female sex. Their taxonomic boundaries can be considered well established, except for Goniozus Förster. Undoubtedly it is the genus with the highest degree of taxonomic confusion regarding its boundaries. The important characters delimiting Goniozus are also shared by several Bethylinae genera, making classification uncertain and hampering understanding of character evolution and variation between taxa. This subfamily currently comprises approximately 540 species described worldwide classified in eight extant genera with low sexual dimorphism. The phylogeny of Bethylinae lineages has received attention by Sorg in 1988, Polaszek & Krombein in 1994, Terayama in 1995 and De Ploëg & Nel in 2004. In all previous analyses performed there is a basal polytomy among Eupsenella, Lytopsenella and the remaining Bethylinae genera. Here we aimed 1) to review the fossil species of Bethylinae, providing descriptions and illustrations when necessary. In addition, o provide a new key to the fossil species of Bethylinae; 2) to recognize, define and describe Afrobethylus as a new Bethylinae genus, its new species. In addition, to define their diagnostic characteristics, as well as provide a key to species of Afrotropical region based on females; 3) to propose a phylogenetic hypothesis based on morphological data of the genera of Bethylinae drawing especial attention to Eupsenella and Lytopsenella relationships with the other genera of this subfamily; 4) to investigate and discuss the main diagnostic characters of Bethylinae genera. Examined material was provided by several institutions. The descriptions, character list, character matrices for cladistic analysis and key, when necessary, were elaborated with o software DELTA. The searches for the most parsimonious trees were carried out under the software TNT. Here, based on our results, we considered that the fossil genus Protobethylus De Ploëg & Nel, 2004 is a junior synonymous of Eupsenella Westwood, 1874. In addition, based on our rescue and compilation of whole the existing information in Bethylinae the fossil family Fushunochrysidae proposed by Hong in 2002 was synonymized with Bethylidae Haliday, 1839. The single genus of this family is Fushunochrysites Hong, 2002 and its single species F. eocenicus Hong, 2002 was established as its type-species. Here, we propose that the best placement of this monotypic genus in Bethylidae is into Bethylinae. Fushunochrysites displays on its forewing several characters that are also present in all members of Eupsenella. Moreover, Sinibethylus Hong, 2002 from Chinese Xilutian coal mine is here also synonymized with Eupsenella. Finally, the last synonymy of genus proposed here is related to Messoria that was described originally by Meunier in 1916. We here propose 10 Messoria as a new junior synonym of Goniozus and transfer its single Messoria copalina Meunier, 1916 to Goniozus. In addition, based on an unusual combination of characters never seen before to Bethylinae we describe Afrobethylus as a new Bethylinae genus. This Bethylinae genus is the unique excluviley Afrotropical. In the phylogenetic context, the resulting matrices contain a total of 61 species of Bethylinae terminals as ingroup, with a total of 43 characters were coded for males and a total of 87 species, 44 characters were coded for females. The resampling analyses based on both males and females returned support for a sister-group relationship among all extant Bethylinae genera. All genera were retrieved as lineages in both analyses, except Goniozus that was retrieved as paraphyletic in all analyses performed. Our most remarkable result obtained was the resolution of the existing basal polytomy in Bethylinae between Lytopsenella and Eupsenella with the other extant Bethylinae genera. In all analyses performed, these genera were retrieved by the first time as sister-group each other and separated from the other genera in all analyses performed. Based on our results, we discuss the main diagnostic characters of Bethylinae genera mainly present in the antenna, forewing, and finally by the first time in the male and female genitalia. In special, about the forewings in Bethylinae, we suggest practicable approaches to functional morphology to the venation in the Bethylinae clades. The present study is the most comprehensive cladistic treatment dedicated to the understanding of the evolution of the Bethylinae genera, and the first to consider a global sampling of species of this subfamily. We conclude that the morphological characters unexplored, mainly, of male and female genitalia might offer additional data relevant to more robustly estimate the phylogenetic history of this group. Here, we also promote a first step to direct and encourage future research in Bethylinae using the arrangement of forewing veins, flexion lines and male and female genitalia characters to phylogenetic reconstruction.Os gêneros de Bethylinae são claramente diferentes um do outro. A maioria das espécies Bethylinae é conhecida apenas pelo sexo feminino. Seus limites taxonômicos podem ser considerados bem estabelecidos, exceto para Goniozus Förster. Sem dúvida é o gênero com o mais alto grau de confusão taxonômica em relação a seus limites. Os caracteres importantes que delimitam Goniozus são compartilhados por vários gêneros de Bethylinae tornando a classificação incerta e dificultando a compreensão da evolução do caráter e da variação entre táxons. Atualmente, esta subfamília compreende aproximadamente 540 espécies descritas classificadas no mundo inteiro em oito gêneros existentes com baixo dimorfismo sexual. A filogenia das linhagens Bethylinae tem recebido atenção por Sorg em 1988, Polaszek & Krombein em 1994, Terayama em 1995 e De Ploëg & Nel em 2004. Em todas as análises realizadas há uma politomia basal entre Eupsenella, Lytopsenella e os demais gêneros de Bethylinae. Aqui objetivamos 1) revisar as espécies fósseis de Bethylinae, fornecendo descrições e ilustrações quando necessário. Além disso, fornecer uma chave nova para as espécies fósseis de Bethylinae; 2) reconhecer, definir e descrever Afrobethylus como um gênero novo para Bethylinae, além de suas espécie novas. Além disso, definir suas características diagnósticas, bem como fornecer uma chave para as espécies da região Afrotropical com base em fêmeas; 3) propor uma hipótese filogenética baseada em dados morfológicos dos gêneros de Bethylinae chamando atenção especial para as relações de Eupsenella e Lytopsenella com os outros gêneros desta subfamília; 4) investigar e discutir os principais caracteres diagnósticos dos gêneros Bethylinae. O material examinado foi fornecido por várias instituições. As descrições, lista de caracteres, matrizes de caracteres para análise cladística e chave, quando necessário, foram elaboradas com o software DELTA. As buscas para as árvores mais parcimoniosas foram realizadas com o software TNT. Aqui, com base nos nossos resultados, consideramos que o gênero fóssil Protobethylus De Ploëg & Nel, 2004 é um sinônimo júnior de Eupsenella Westwood, 1874. Além disso, com base em nosso resgate e compilação de toda a informação existente em Bethylinae a família fóssil Fushunochrysidae proposta por Hong em 2002 foi sinonimizada com Bethylidae Haliday, 1839. O único gênero desta família é Fushunochrysites Hong, 2002 e sua única espécie F. eocenicus Hong, 2002 foi estabelecida como sua espécie tipo. Aqui, propomos que a melhor alocação deste gênero monotípico em Bethylidae é em Bethylinae. Fushunochrysites exibe em sua asa anterior vários caracteres que também estão presentes em todos os membros de Eupsenella. Além disso, Sinibethylus Hong, 2002 da mina de carvão chinesa Xilutian é aqui 8 também sinonimizada com Eupsenella. Finalmente, a última sinonímia do gênero proposta aqui está relacionada à Messoria que foi descrita originalmente por Meunier em 1916. Aqui propomos Messoria como sinônimo júnior de Goniozus e transferir sua única espécie Messoria copalina Meunier, 1916 para Goniozus. Além disso, com base em uma incomum combinação de caracteres descrevemos Afrobethylus como um novo gênero Bethylinae. Este gênero de Bethylinae é o único exclusivamente Afrotropical. No contexto filogenético, as matrizes resultantes contêm um total de 61 espécies terminais de Bethylinae como grupo interno, com um total de 43 caracteres codificados para machos e um total de 87 espécies, 44 caracteres codificados para fêmeas. As análises de reamostragem baseadas em machos e fêmeas retornaram suporte para uma relação de grupo irmão entre todos os gêneros de Bethylinae existentes. Todos os gêneros foram recuperados como linhagens em ambas as análises, exceto Goniozus que foi recuperado como parafilético em todas as análises realizadas. Nosso mais notável resultado obtido foi à resolução da politomia basal existente em Bethylinae entre Lytopsenella e Eupsenella com os outros gêneros Bethylinae existentes. Em todas as análises realizadas, esses gêneros foram recuperados pela primeira vez como um grupo irmão entre si e separados dos demais gêneros em todas as análises realizadas. Com base em nossos resultados, discutimos os principais caracteres diagnósticos dos gêneros Bethylinae, presentes principalmente na antena, na asa anterior e, finalmente, pela primeira vez nas genitálias dos machos e das fêmeas. Em especial, sobre as asas anteriores em Bethylinae, sugerimos abordagens viáveis para à morfologia funcional da venação nos clados de Bethylinae. O presente estudo é o tratamento cladístico mais abrangente dedicado à compreensão da evolução dos gêneros de Bethylinae e o primeiro a considerar uma amostragem global das espécies desta subfamília. Concluímos que os caracteres morfológicos inexplorados, principalmente, das genitálias dos machos e das fêmeas podem oferecer dados adicionais relevantes para estimar de forma mais robusta a história filogenética deste grupo. Aqui, promovemos um primeiro passo para direcionar e encorajar futuras pesquisas em Bethylinae usando o arranjo de veias, linhas de flexão e caracteres da genitália dos machos e das fêmeas para a reconstrução filogenética.TextRAMOS, Magno Suprani. Sistemática de Bethylinae (Hymenoptera, Bethylidae). 2017. 195 f. Tese (Doutorado em Biologia Animal) - Universidade Federal do Espírito Santo, Centro de Ciências Humanas e Naturais, Vitória, 2017.http://repositorio.ufes.br/handle/10/9917porUniversidade Federal do Espírito SantoDoutorado em Biologia AnimalPrograma de Pós-Graduação em Ciências BiológicasUFESBRBethylidaeZoologia x ClassificaçãoInsetoFilogeniaHimenópteroZoologia57Sistemática de Bethylinae (Hymenoptera, Bethylidae)info:eu-repo/semantics/publishedVersioninfo:eu-repo/semantics/doctoralThesisinfo:eu-repo/semantics/openAccessreponame:Repositório Institucional da Universidade Federal do Espírito Santo (riUfes)instname:Universidade Federal do Espírito Santo (UFES)instacron:UFESORIGINALTese Magno _ Versão Final.pdfapplication/pdf7825784http://repositorio.ufes.br/bitstreams/8bcfc975-0d01-4968-a59e-95c5a3f186d4/download72c21f52127847746a1c8973b4ff84ecMD5110/99172024-07-01 16:23:47.341oai:repositorio.ufes.br:10/9917http://repositorio.ufes.brRepositório InstitucionalPUBhttp://repositorio.ufes.br/oai/requestopendoar:21082024-07-11T14:26:44.740436Repositório Institucional da Universidade Federal do Espírito Santo (riUfes) - Universidade Federal do Espírito Santo (UFES)false |
dc.title.none.fl_str_mv |
Sistemática de Bethylinae (Hymenoptera, Bethylidae) |
title |
Sistemática de Bethylinae (Hymenoptera, Bethylidae) |
spellingShingle |
Sistemática de Bethylinae (Hymenoptera, Bethylidae) Ramos, Magno Suprani Zoologia Bethylidae Zoologia x Classificação Inseto Filogenia Himenóptero 57 |
title_short |
Sistemática de Bethylinae (Hymenoptera, Bethylidae) |
title_full |
Sistemática de Bethylinae (Hymenoptera, Bethylidae) |
title_fullStr |
Sistemática de Bethylinae (Hymenoptera, Bethylidae) |
title_full_unstemmed |
Sistemática de Bethylinae (Hymenoptera, Bethylidae) |
title_sort |
Sistemática de Bethylinae (Hymenoptera, Bethylidae) |
author |
Ramos, Magno Suprani |
author_facet |
Ramos, Magno Suprani |
author_role |
author |
dc.contributor.advisor1.fl_str_mv |
Azevedo, Celso Oliveira |
dc.contributor.author.fl_str_mv |
Ramos, Magno Suprani |
dc.contributor.referee1.fl_str_mv |
Aguiar, Alexandre Pires |
dc.contributor.referee2.fl_str_mv |
Monteiro, Cecília Waichert |
dc.contributor.referee3.fl_str_mv |
Gobbi, Fernanda Tonini |
dc.contributor.referee4.fl_str_mv |
Hermes, Marcel Gustavo |
contributor_str_mv |
Azevedo, Celso Oliveira Aguiar, Alexandre Pires Monteiro, Cecília Waichert Gobbi, Fernanda Tonini Hermes, Marcel Gustavo |
dc.subject.cnpq.fl_str_mv |
Zoologia |
topic |
Zoologia Bethylidae Zoologia x Classificação Inseto Filogenia Himenóptero 57 |
dc.subject.br-rjbn.none.fl_str_mv |
Bethylidae Zoologia x Classificação Inseto Filogenia Himenóptero |
dc.subject.udc.none.fl_str_mv |
57 |
description |
The genera of Bethylinae are clearly different one from another. The most of Bethylinae species are known only by the female sex. Their taxonomic boundaries can be considered well established, except for Goniozus Förster. Undoubtedly it is the genus with the highest degree of taxonomic confusion regarding its boundaries. The important characters delimiting Goniozus are also shared by several Bethylinae genera, making classification uncertain and hampering understanding of character evolution and variation between taxa. This subfamily currently comprises approximately 540 species described worldwide classified in eight extant genera with low sexual dimorphism. The phylogeny of Bethylinae lineages has received attention by Sorg in 1988, Polaszek & Krombein in 1994, Terayama in 1995 and De Ploëg & Nel in 2004. In all previous analyses performed there is a basal polytomy among Eupsenella, Lytopsenella and the remaining Bethylinae genera. Here we aimed 1) to review the fossil species of Bethylinae, providing descriptions and illustrations when necessary. In addition, o provide a new key to the fossil species of Bethylinae; 2) to recognize, define and describe Afrobethylus as a new Bethylinae genus, its new species. In addition, to define their diagnostic characteristics, as well as provide a key to species of Afrotropical region based on females; 3) to propose a phylogenetic hypothesis based on morphological data of the genera of Bethylinae drawing especial attention to Eupsenella and Lytopsenella relationships with the other genera of this subfamily; 4) to investigate and discuss the main diagnostic characters of Bethylinae genera. Examined material was provided by several institutions. The descriptions, character list, character matrices for cladistic analysis and key, when necessary, were elaborated with o software DELTA. The searches for the most parsimonious trees were carried out under the software TNT. Here, based on our results, we considered that the fossil genus Protobethylus De Ploëg & Nel, 2004 is a junior synonymous of Eupsenella Westwood, 1874. In addition, based on our rescue and compilation of whole the existing information in Bethylinae the fossil family Fushunochrysidae proposed by Hong in 2002 was synonymized with Bethylidae Haliday, 1839. The single genus of this family is Fushunochrysites Hong, 2002 and its single species F. eocenicus Hong, 2002 was established as its type-species. Here, we propose that the best placement of this monotypic genus in Bethylidae is into Bethylinae. Fushunochrysites displays on its forewing several characters that are also present in all members of Eupsenella. Moreover, Sinibethylus Hong, 2002 from Chinese Xilutian coal mine is here also synonymized with Eupsenella. Finally, the last synonymy of genus proposed here is related to Messoria that was described originally by Meunier in 1916. We here propose 10 Messoria as a new junior synonym of Goniozus and transfer its single Messoria copalina Meunier, 1916 to Goniozus. In addition, based on an unusual combination of characters never seen before to Bethylinae we describe Afrobethylus as a new Bethylinae genus. This Bethylinae genus is the unique excluviley Afrotropical. In the phylogenetic context, the resulting matrices contain a total of 61 species of Bethylinae terminals as ingroup, with a total of 43 characters were coded for males and a total of 87 species, 44 characters were coded for females. The resampling analyses based on both males and females returned support for a sister-group relationship among all extant Bethylinae genera. All genera were retrieved as lineages in both analyses, except Goniozus that was retrieved as paraphyletic in all analyses performed. Our most remarkable result obtained was the resolution of the existing basal polytomy in Bethylinae between Lytopsenella and Eupsenella with the other extant Bethylinae genera. In all analyses performed, these genera were retrieved by the first time as sister-group each other and separated from the other genera in all analyses performed. Based on our results, we discuss the main diagnostic characters of Bethylinae genera mainly present in the antenna, forewing, and finally by the first time in the male and female genitalia. In special, about the forewings in Bethylinae, we suggest practicable approaches to functional morphology to the venation in the Bethylinae clades. The present study is the most comprehensive cladistic treatment dedicated to the understanding of the evolution of the Bethylinae genera, and the first to consider a global sampling of species of this subfamily. We conclude that the morphological characters unexplored, mainly, of male and female genitalia might offer additional data relevant to more robustly estimate the phylogenetic history of this group. Here, we also promote a first step to direct and encourage future research in Bethylinae using the arrangement of forewing veins, flexion lines and male and female genitalia characters to phylogenetic reconstruction. |
publishDate |
2017 |
dc.date.issued.fl_str_mv |
2017-02-01 |
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info:eu-repo/semantics/publishedVersion |
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info:eu-repo/semantics/doctoralThesis |
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doctoralThesis |
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publishedVersion |
dc.identifier.citation.fl_str_mv |
RAMOS, Magno Suprani. Sistemática de Bethylinae (Hymenoptera, Bethylidae). 2017. 195 f. Tese (Doutorado em Biologia Animal) - Universidade Federal do Espírito Santo, Centro de Ciências Humanas e Naturais, Vitória, 2017. |
dc.identifier.uri.fl_str_mv |
http://repositorio.ufes.br/handle/10/9917 |
identifier_str_mv |
RAMOS, Magno Suprani. Sistemática de Bethylinae (Hymenoptera, Bethylidae). 2017. 195 f. Tese (Doutorado em Biologia Animal) - Universidade Federal do Espírito Santo, Centro de Ciências Humanas e Naturais, Vitória, 2017. |
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http://repositorio.ufes.br/handle/10/9917 |
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Text |
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Universidade Federal do Espírito Santo Doutorado em Biologia Animal |
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Programa de Pós-Graduação em Ciências Biológicas |
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UFES |
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BR |
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Universidade Federal do Espírito Santo Doutorado em Biologia Animal |
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