Análise filogenética de Gymnophiona Müller, 1832 (Amphibia)
Autor(a) principal: | |
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Data de Publicação: | 2016 |
Tipo de documento: | Dissertação |
Idioma: | por |
Título da fonte: | Biblioteca Digital de Teses e Dissertações da PUC_RS |
Texto Completo: | http://tede2.pucrs.br/tede2/handle/tede/7723 |
Resumo: | Gymnophiona Müller, 1832—with 206 currently recognized species—is the least studied order within Amphibia. The phylogenetic relationships among its members have been historically unstable, with frequent taxonomic changes at the family level due to the recurrent presence of paraphyletic taxa. However, an increase of studies based on morphological and molecular data has built a scaffold of information about the evolutionary relationships among caecilians; even though, a phylogenetic classification of the order has remained a challenge. In 2011 a new taxonomy of Gymnophiona was proposed, where genera were arranged in nine families supposedly monophyletic. This taxonomy was not based on a phylogenetic analysis but on a consensus of the understanding of the evolutionary relationships of the group as inferred by previous studies. However, important conflicts exits among the results and type of analyses performed on these studies. Furthermore, only one study has included the known fossil taxon of Gymnophiona, which despite being fragmented is a valid source of evidence. Due to several and continuous efforts to generate data on caecilians, there is a wealth of hereditary characters available for phylogenetic studies. Thus, it has become crucial to perform a combined analysis of all these data (complete mitochondrial genomes, nuclear DNA sequences, phenotypic characters of extant and fossil taxa) to obtain a phylogenetic hypothesis that maximizes explanatory power. The objective of this study is to infer the evolutionary relationships of Gymnophiona based on a total evidence analysis using parsimony and dynamic homology to evaluate the current taxonomy of the group. Additionally, we also evaluate the effect of different types of alignment (tree versus similarity approaches), the influence of phenotypic characters on a dataset dominated by molecular characters, and the effect of coding indels as missing data. We compiled previously published phenotypic characters, DNA sequences available from GenBank of 47 nuclear and mitochondrial genes of all available taxa, and produced 42 new sequences. When comparing the results obtained from the tree-alignment analysis in POY with those of the similarity alignment in TNT, both recover Rhinatrematidae as sister of Ichthyophiidae + Teresomata. Within Rhinatrematidae, Epicrionops is paraphyletic in all analyses and the relationships within Ichthyophis are unsolved (a polytomy in the strict consensus). POY does not recovered Scolecomorphidae as the sister taxon of all other Teresomata but Typhlonectidae + Caeciliidae. Caecilia is paraphyletic with respect to Oscaecilia and Typhlonectes in relationship to Potamotyphlus. Scolecomorphidae is sister of Herpelidae + Chikilidae. The results of TNT recover a paraphyletic Herpelidae, with Herpele squalostoma sister of Chikilidae. Also, Siphonopidae is non-monophyletic. Indotyphlidae is non-monophyletic in both analyses and Idiocranium is consistently recovered as sister taxon of Dermophiidae. Dermophis is recovered as paraphyletic in the POY analysis, while both Dermophis and Gymnopis are paraphyletic in the TNT analysis. The strict consensus of the molecular dataset is highly congruous with that of the total evidence dataset; however, the former is better resolved (less polytomies), mainly within Indotyphlidae. The analysis of the phenotypic data alone resulted in a complete polytomy, illustrating the need of more research in this avenue. Coding indels as missing data did not cause important topological changes. The main conclusions derived from this study are: (i) the type of alignment of DNA sequences have an evident impact on the phylogenetic hypotheses of Gymnophiona; (ii) the apparent resolution on the evolutionary relationships of the extant supraspecific taxa of Gymnophiona and their monophyly presented in other studies are dependent on the exclusion of relevant evidence—taxa and characters—or the partial presentation of the optimal hypotheses; (iii) only a total evidence analysis allowed us to discover some of the potential cases of paraphyly or misidentification of vouchers; (iv) the phenotypic data currently used in the study of the evolutionary relationships of Gymnophiona contain important levels of non-congruent information and are not sufficient to place the fossil Eocaecilia micropodia within caecilians. This study reveals the need of detailed revision of the taxonomy and phylogeny of Gymnophiona. |
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Castroviejo-Fisher, Santiagohttp://buscatextual.cnpq.br/buscatextual/visualizacv.do?id=K4802148P9http://buscatextual.cnpq.br/buscatextual/visualizacv.do?id=K4337415Y3Souza, Camila Camargo de2017-11-09T17:46:23Z2016-03-30http://tede2.pucrs.br/tede2/handle/tede/7723Gymnophiona Müller, 1832—with 206 currently recognized species—is the least studied order within Amphibia. The phylogenetic relationships among its members have been historically unstable, with frequent taxonomic changes at the family level due to the recurrent presence of paraphyletic taxa. However, an increase of studies based on morphological and molecular data has built a scaffold of information about the evolutionary relationships among caecilians; even though, a phylogenetic classification of the order has remained a challenge. In 2011 a new taxonomy of Gymnophiona was proposed, where genera were arranged in nine families supposedly monophyletic. This taxonomy was not based on a phylogenetic analysis but on a consensus of the understanding of the evolutionary relationships of the group as inferred by previous studies. However, important conflicts exits among the results and type of analyses performed on these studies. Furthermore, only one study has included the known fossil taxon of Gymnophiona, which despite being fragmented is a valid source of evidence. Due to several and continuous efforts to generate data on caecilians, there is a wealth of hereditary characters available for phylogenetic studies. Thus, it has become crucial to perform a combined analysis of all these data (complete mitochondrial genomes, nuclear DNA sequences, phenotypic characters of extant and fossil taxa) to obtain a phylogenetic hypothesis that maximizes explanatory power. The objective of this study is to infer the evolutionary relationships of Gymnophiona based on a total evidence analysis using parsimony and dynamic homology to evaluate the current taxonomy of the group. Additionally, we also evaluate the effect of different types of alignment (tree versus similarity approaches), the influence of phenotypic characters on a dataset dominated by molecular characters, and the effect of coding indels as missing data. We compiled previously published phenotypic characters, DNA sequences available from GenBank of 47 nuclear and mitochondrial genes of all available taxa, and produced 42 new sequences. When comparing the results obtained from the tree-alignment analysis in POY with those of the similarity alignment in TNT, both recover Rhinatrematidae as sister of Ichthyophiidae + Teresomata. Within Rhinatrematidae, Epicrionops is paraphyletic in all analyses and the relationships within Ichthyophis are unsolved (a polytomy in the strict consensus). POY does not recovered Scolecomorphidae as the sister taxon of all other Teresomata but Typhlonectidae + Caeciliidae. Caecilia is paraphyletic with respect to Oscaecilia and Typhlonectes in relationship to Potamotyphlus. Scolecomorphidae is sister of Herpelidae + Chikilidae. The results of TNT recover a paraphyletic Herpelidae, with Herpele squalostoma sister of Chikilidae. Also, Siphonopidae is non-monophyletic. Indotyphlidae is non-monophyletic in both analyses and Idiocranium is consistently recovered as sister taxon of Dermophiidae. Dermophis is recovered as paraphyletic in the POY analysis, while both Dermophis and Gymnopis are paraphyletic in the TNT analysis. The strict consensus of the molecular dataset is highly congruous with that of the total evidence dataset; however, the former is better resolved (less polytomies), mainly within Indotyphlidae. The analysis of the phenotypic data alone resulted in a complete polytomy, illustrating the need of more research in this avenue. Coding indels as missing data did not cause important topological changes. The main conclusions derived from this study are: (i) the type of alignment of DNA sequences have an evident impact on the phylogenetic hypotheses of Gymnophiona; (ii) the apparent resolution on the evolutionary relationships of the extant supraspecific taxa of Gymnophiona and their monophyly presented in other studies are dependent on the exclusion of relevant evidence—taxa and characters—or the partial presentation of the optimal hypotheses; (iii) only a total evidence analysis allowed us to discover some of the potential cases of paraphyly or misidentification of vouchers; (iv) the phenotypic data currently used in the study of the evolutionary relationships of Gymnophiona contain important levels of non-congruent information and are not sufficient to place the fossil Eocaecilia micropodia within caecilians. This study reveals the need of detailed revision of the taxonomy and phylogeny of Gymnophiona.Gymnophiona Müller, 1832 é a ordem menos estudada dentro de Amphibia, com 206 espécies conhecidas. As relações filogenéticas do grupo permaneceram instáveis por muito tempo, sofrendo com mudanças no número de famílias, devido à constante presença de táxons parafiléticos. Uma crescente de estudos com produção de evidências moleculares e morfológicas tem formado um arcabouço de informações sobre as relações evolutivas do grupo. Mesmo assim, uma classificação filogenética de toda a ordem não representava uma tarefa fácil. Em 2011, foi proposta uma classificação ao nível de família, onde os gêneros foram distribuídos em nove famílias supostamente monofiléticas. Essa proposta não foi baseada em uma análise filogenética, mas sim em um consenso do entendimento das relações filogenéticas inferidas em estudos prévios, sendo muitos deles discordantes, tanto em tipos de análises, quanto em resultados. Apenas um estudo incluiu um táxon fóssil, que apesar de ser pouco representativo em quantidade, representa uma fonte válida de evidência. Devido ao grande esforço para gerar dados e obter informações acerca dos Gymnophiona, existem inúmeros caracteres hereditários disponíveis. Dessa forma, percebe-se uma importância em unir todos esses dados (genomas mitocondriais completos, sequências de genes nucleares, caracteres fenotípicos dos táxons viventes e fóssil) para compor uma nova proposta filogenética com o maior poder explicativo. Assim, o objetivo principal deste estudo foi inferir as relações evolutivas da ordem Gymnophiona em um contexto de evidência total, sob análises de parcimônia e homologia dinâmica e avaliar a taxonomia atual de Gymnophiona com base na(s) árvore(s) mais parcimoniosa(s). Adicionalmente, avaliar o efeito de diferentes tipos de alinhamento, a influência de caracteres fenotípicos em um dataset dominado por dados genéticos e o efeito de codificar os indels como dados faltantes. Para isso foram compilados caracteres morfológicos de estudos prévios, sequências disponíveis no GenBank, referentes a 47 genes (nucleares e mitocondriais) de todas as espécies disponíveis, além da produção de 42 novas sequências. Comparando os resultados gerados a partir do alinhamento por árvore (POY) e do alinhamento por similaridade (TNT) encontramos Rhinatrematidae como grupo-irmão de Ichthyophiidae + Teresomata. Dentro de Rhinatrematidae, Epicrionops foi recuperado parafilético em todas as análises e as relações dentro de Ichthyophiidae são uma politomia. A análise do POY não recupera Scolecomorphidae como táxon-irmão dos demais Teresomata, e sim Typhlonectidae + Caeciliidae. Caecilia é parafilética a Oscaecilia, assim como Typhlonectes com relação a Potamotyphlus. Na análise via TNT, Herpelidae foi recuperada parafilética, com Herpele squalostoma agrupado com Chikilidae. Na análise do TNT, Siphonopidae não foi recuperada monofilética. A família Indotyphlidae foi não-monofilética em ambas as análises e Idiocranium foi recuperado como o táxon-irmão de Dermophiidae. Dermophis surge parafilético na análise do POY, enquanto que pelo TNT tanto Dermophis quanto Gymnopis surgen parafiléticos. A topologia da árvore construída apenas com sequências de DNA, mostrou-se altamente semelhante ao consenso gerado pelo TNT, embora com alguns poucos clados melhor resolvidos, principalmente em Indotyphlidae. Já a análise unicamente morfológica restringiu-se a uma politomia total, o que pode ser justificado pela falta de caracteres morfológicos úteis, e que também pode ser um reflexo da falta de conhecimento acerca deste tipo de evidência. Quanto à codificação de indels como quinto estado ou como dados faltantes, não foi observada vantagem sobre nenhum dos testes em relação à topologia do consenso estrito. As principais conclusões derivadas dos resultados obtidos são: (i) o tipo de alinhamento das sequências de DNA tem um impacto evidente nas hipóteses filogenéticas; (ii) a aparente resolução da história evolutiva dos atuais táxons supraespecíficos de Gymnophiona e sua monofilia, apresentada em outros estudos, dependem da exclusão de evidência relevante das análises ou da apresentação parcial das hipóteses ótimas; (iii) só a análise de evidência total permitiu descobrir casos potenciais de parafilia ou identificação errada de terminais; (iv) os dados fenotípicos atualmente usados no estudo das relações evolutivas de Gymnophiona têm altos níveis de informação não congruente e não são suficentes para a inclusão do táxon fóssil Eocaecilia micropodia dentro de Gymnophiona. O presente estudo revela a necessidade de uma revisão detalhada da taxonomia e filogenética de Gymnophiona.Submitted by Caroline Xavier (caroline.xavier@pucrs.br) on 2017-11-09T17:46:02Z No. of bitstreams: 1 DIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf: 3948791 bytes, checksum: c827f34181e47b84d2dc6c25fa596df9 (MD5)Approved for entry into archive by Caroline Xavier (caroline.xavier@pucrs.br) on 2017-11-09T17:46:13Z (GMT) No. of bitstreams: 1 DIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf: 3948791 bytes, checksum: c827f34181e47b84d2dc6c25fa596df9 (MD5)Made available in DSpace on 2017-11-09T17:46:23Z (GMT). No. of bitstreams: 1 DIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf: 3948791 bytes, checksum: c827f34181e47b84d2dc6c25fa596df9 (MD5) Previous issue date: 2016-03-30Conselho Nacional de Pesquisa e Desenvolvimento Científico e Tecnológico - CNPqapplication/pdfhttp://tede2.pucrs.br:80/tede2/retrieve/170243/DIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf.jpgporPontifícia Universidade Católica do Rio Grande do SulPrograma de Pós Graduação em ZoologiaPUCRSBrasilFaculdade de BiociênciasCecíliasSistemática FilogenéticaParcimôniaAlinhamento por ÁrvoreAlinhamento por SimilaridadeEvidência TotalCIENCIAS BIOLOGICAS::ZOOLOGIAAnálise filogenética de Gymnophiona Müller, 1832 (Amphibia)info:eu-repo/semantics/publishedVersioninfo:eu-repo/semantics/masterThesisTrabalho não apresenta restrição para publicação36528317262667714500500600-6482652380601267558-2555911436985713659info:eu-repo/semantics/openAccessreponame:Biblioteca Digital de Teses e Dissertações da PUC_RSinstname:Pontifícia Universidade Católica do Rio Grande do Sul (PUCRS)instacron:PUC_RSTHUMBNAILDIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf.jpgDIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf.jpgimage/jpeg3971http://tede2.pucrs.br/tede2/bitstream/tede/7723/4/DIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf.jpgbe3be406db731b321c2d972ef60db636MD54TEXTDIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf.txtDIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf.txttext/plain126112http://tede2.pucrs.br/tede2/bitstream/tede/7723/3/DIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdf.txt31cddec9bf5bfcc513b3cb430a948d3fMD53ORIGINALDIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdfDIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdfapplication/pdf3948791http://tede2.pucrs.br/tede2/bitstream/tede/7723/2/DIS_CAMILA_CAMARGO_DE_SOUZA_COMPLETO.pdfc827f34181e47b84d2dc6c25fa596df9MD52LICENSElicense.txtlicense.txttext/plain; charset=utf-8610http://tede2.pucrs.br/tede2/bitstream/tede/7723/1/license.txt5a9d6006225b368ef605ba16b4f6d1beMD51tede/77232017-11-09 20:00:56.393oai:tede2.pucrs.br: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Biblioteca Digital de Teses e Dissertaçõeshttp://tede2.pucrs.br/tede2/PRIhttps://tede2.pucrs.br/oai/requestbiblioteca.central@pucrs.br||opendoar:2017-11-09T22:00:56Biblioteca Digital de Teses e Dissertações da PUC_RS - Pontifícia Universidade Católica do Rio Grande do Sul (PUCRS)false |
dc.title.por.fl_str_mv |
Análise filogenética de Gymnophiona Müller, 1832 (Amphibia) |
title |
Análise filogenética de Gymnophiona Müller, 1832 (Amphibia) |
spellingShingle |
Análise filogenética de Gymnophiona Müller, 1832 (Amphibia) Souza, Camila Camargo de Cecílias Sistemática Filogenética Parcimônia Alinhamento por Árvore Alinhamento por Similaridade Evidência Total CIENCIAS BIOLOGICAS::ZOOLOGIA |
title_short |
Análise filogenética de Gymnophiona Müller, 1832 (Amphibia) |
title_full |
Análise filogenética de Gymnophiona Müller, 1832 (Amphibia) |
title_fullStr |
Análise filogenética de Gymnophiona Müller, 1832 (Amphibia) |
title_full_unstemmed |
Análise filogenética de Gymnophiona Müller, 1832 (Amphibia) |
title_sort |
Análise filogenética de Gymnophiona Müller, 1832 (Amphibia) |
author |
Souza, Camila Camargo de |
author_facet |
Souza, Camila Camargo de |
author_role |
author |
dc.contributor.advisor1.fl_str_mv |
Castroviejo-Fisher, Santiago |
dc.contributor.advisor1Lattes.fl_str_mv |
http://buscatextual.cnpq.br/buscatextual/visualizacv.do?id=K4802148P9 |
dc.contributor.authorLattes.fl_str_mv |
http://buscatextual.cnpq.br/buscatextual/visualizacv.do?id=K4337415Y3 |
dc.contributor.author.fl_str_mv |
Souza, Camila Camargo de |
contributor_str_mv |
Castroviejo-Fisher, Santiago |
dc.subject.por.fl_str_mv |
Cecílias Sistemática Filogenética Parcimônia Alinhamento por Árvore Alinhamento por Similaridade Evidência Total |
topic |
Cecílias Sistemática Filogenética Parcimônia Alinhamento por Árvore Alinhamento por Similaridade Evidência Total CIENCIAS BIOLOGICAS::ZOOLOGIA |
dc.subject.cnpq.fl_str_mv |
CIENCIAS BIOLOGICAS::ZOOLOGIA |
description |
Gymnophiona Müller, 1832—with 206 currently recognized species—is the least studied order within Amphibia. The phylogenetic relationships among its members have been historically unstable, with frequent taxonomic changes at the family level due to the recurrent presence of paraphyletic taxa. However, an increase of studies based on morphological and molecular data has built a scaffold of information about the evolutionary relationships among caecilians; even though, a phylogenetic classification of the order has remained a challenge. In 2011 a new taxonomy of Gymnophiona was proposed, where genera were arranged in nine families supposedly monophyletic. This taxonomy was not based on a phylogenetic analysis but on a consensus of the understanding of the evolutionary relationships of the group as inferred by previous studies. However, important conflicts exits among the results and type of analyses performed on these studies. Furthermore, only one study has included the known fossil taxon of Gymnophiona, which despite being fragmented is a valid source of evidence. Due to several and continuous efforts to generate data on caecilians, there is a wealth of hereditary characters available for phylogenetic studies. Thus, it has become crucial to perform a combined analysis of all these data (complete mitochondrial genomes, nuclear DNA sequences, phenotypic characters of extant and fossil taxa) to obtain a phylogenetic hypothesis that maximizes explanatory power. The objective of this study is to infer the evolutionary relationships of Gymnophiona based on a total evidence analysis using parsimony and dynamic homology to evaluate the current taxonomy of the group. Additionally, we also evaluate the effect of different types of alignment (tree versus similarity approaches), the influence of phenotypic characters on a dataset dominated by molecular characters, and the effect of coding indels as missing data. We compiled previously published phenotypic characters, DNA sequences available from GenBank of 47 nuclear and mitochondrial genes of all available taxa, and produced 42 new sequences. When comparing the results obtained from the tree-alignment analysis in POY with those of the similarity alignment in TNT, both recover Rhinatrematidae as sister of Ichthyophiidae + Teresomata. Within Rhinatrematidae, Epicrionops is paraphyletic in all analyses and the relationships within Ichthyophis are unsolved (a polytomy in the strict consensus). POY does not recovered Scolecomorphidae as the sister taxon of all other Teresomata but Typhlonectidae + Caeciliidae. Caecilia is paraphyletic with respect to Oscaecilia and Typhlonectes in relationship to Potamotyphlus. Scolecomorphidae is sister of Herpelidae + Chikilidae. The results of TNT recover a paraphyletic Herpelidae, with Herpele squalostoma sister of Chikilidae. Also, Siphonopidae is non-monophyletic. Indotyphlidae is non-monophyletic in both analyses and Idiocranium is consistently recovered as sister taxon of Dermophiidae. Dermophis is recovered as paraphyletic in the POY analysis, while both Dermophis and Gymnopis are paraphyletic in the TNT analysis. The strict consensus of the molecular dataset is highly congruous with that of the total evidence dataset; however, the former is better resolved (less polytomies), mainly within Indotyphlidae. The analysis of the phenotypic data alone resulted in a complete polytomy, illustrating the need of more research in this avenue. Coding indels as missing data did not cause important topological changes. The main conclusions derived from this study are: (i) the type of alignment of DNA sequences have an evident impact on the phylogenetic hypotheses of Gymnophiona; (ii) the apparent resolution on the evolutionary relationships of the extant supraspecific taxa of Gymnophiona and their monophyly presented in other studies are dependent on the exclusion of relevant evidence—taxa and characters—or the partial presentation of the optimal hypotheses; (iii) only a total evidence analysis allowed us to discover some of the potential cases of paraphyly or misidentification of vouchers; (iv) the phenotypic data currently used in the study of the evolutionary relationships of Gymnophiona contain important levels of non-congruent information and are not sufficient to place the fossil Eocaecilia micropodia within caecilians. This study reveals the need of detailed revision of the taxonomy and phylogeny of Gymnophiona. |
publishDate |
2016 |
dc.date.issued.fl_str_mv |
2016-03-30 |
dc.date.accessioned.fl_str_mv |
2017-11-09T17:46:23Z |
dc.type.status.fl_str_mv |
info:eu-repo/semantics/publishedVersion |
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http://tede2.pucrs.br/tede2/handle/tede/7723 |
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http://tede2.pucrs.br/tede2/handle/tede/7723 |
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Pontifícia Universidade Católica do Rio Grande do Sul |
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Programa de Pós Graduação em Zoologia |
dc.publisher.initials.fl_str_mv |
PUCRS |
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Brasil |
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Faculdade de Biociências |
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Pontifícia Universidade Católica do Rio Grande do Sul |
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