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AVIS Ce Document a Été Numérisé Par La Division De La Gestion Des Direction des bibliothèques AVIS Ce document a été numérisé par la Division de la gestion des documents et des archives de l’Université de Montréal. L’auteur a autorisé l’Université de Montréal à reproduire et diffuser, en totalité ou en partie, par quelque moyen que ce soit et sur quelque support que ce soit, et exclusivement à des fins non lucratives d’enseignement et de recherche, des copies de ce mémoire ou de cette thèse. L’auteur et les coauteurs le cas échéant conservent la propriété du droit d’auteur et des droits moraux qui protègent ce document. Ni la thèse ou le mémoire, ni des extraits substantiels de ce document, ne doivent être imprimés ou autrement reproduits sans l’autorisation de l’auteur. Afin de se conformer à la Loi canadienne sur la protection des renseignements personnels, quelques formulaires secondaires, coordonnées ou signatures intégrées au texte ont pu être enlevés de ce document. Bien que cela ait pu affecter la pagination, il n’y a aucun contenu manquant. NOTICE This document was digitized by the Records Management & Archives Division of Université de Montréal. The author of this thesis or dissertation has granted a nonexclusive license allowing Université de Montréal to reproduce and publish the document, in part or in whole, and in any format, solely for noncommercial educational and research purposes. The author and co-authors if applicable retain copyright ownership and moral rights in this document. Neither the whole thesis or dissertation, nor substantial extracts from it, may be printed or otherwise reproduced without the author’s permission. In compliance with the Canadian Privacy Act some supporting forms, contact information or signatures may have been removed from the document. While this may affect the document page count, it does not represent any loss of content from the document. Université de Montréal Origin of Symphyotricltum anticostense (Asteraceae: Astereae), an endemic species of the Gulf of St. Lawrence par Jami! Vaezi Département de Sciences biologiques Université de Montréal Thèse présentée à la Faculté des études supérieures en vue de l'obtention du grade de Ph.D. en Sciences biologiques Juillet 2008 © Jamit Vaezi, 2008 Université de Montréal Faculté des etudes supérieures Cette the se intitulée: Origin of Symphyotrichum altticosteltse (Asteraceae: Astereae), an endemic species of the Gulf of St. Lawrence présentée par: Jami! Vaezi a été évaluée par un jury composé des personnes suivantes: Anne Bruneau, Présidente rapporteuse Luc Brouillet, directeur de recherche Mohamed Rijri, membre du jury Geraldine A. Allen, examinatrice externe Représentant du doyen de la FES ii Abstract This thesis discusses the origin of Symphyotrichum anticostense, a decaploid species restricted to the Gulf of St. Lawrence region (Quebec, New Brunswick, and Maine). Two closely related species, S. novi-belgii, a hexaploid and S. boreale, a putatively tetraploid species have been proposed to be putative parents of S. anticostense. Symphyotrichum novi-belgii is distributed in the eastern coastal region of North America and S. boreale is widespread throughout the northern part of North America from west to east. These three species belong to the genus Symphyotrichum, which comprises ca. 91 species ofwhich more th an 50% are polyploids. The evolutionary history of the genus is poorly known even at the diploidy level. Before investigating the origin of S. anticostense, it seems necessary in the first step to elucidate phylogenetic relationships within the genus. The objectives ofthis thesis are therefore to delimit species boundaries excluding polyploid species and to investigate the origin of S. anticostense. The ITS data set nearly supports previous classifications based on morphological and cytological approaches, while the GAPDH nuclear gene is in poor agreement with the ITS results. Incomplete lineage sorting within Symphyotrichum is shown by stochastic sorting of ancestral polymorphisms among the GAPDH alleles. This evidence, in combination with the insufficient resolution among ribotypes, suggests recent evolutionary radiation of the genus. Univariate and multivariate morphological analyses as weil as the ITS results roughly support S. anticostense to be a hybrid derivative of S. novi-belgii and S. boreale. The ITS-based network analysis suggests three independent geographical origins for S. anticostellse: 1) Lake St. John; 2) Gaspé Peninsula- New Brunswick- Maine; 3) Anticosti Island. The results oftwo nuclear genes (GAPDH and MIPS) demonstrate relative incongruence between the phylogenetic trees. Although the results obtained from the 111 latter markers support those obtained from the morphological and ITS data sets, incomplete lineage sorting strongly affected the phylogenetic relationships among species. Key words: Asteraceae, Astereae, Symphyotrichum, Symphyotrichum anticostellse, polyploidy, hybridization, phylogeny, incongruence, incomplete lineage sorting, radiation. IV Résumé Cette thèse porte sur l'origine de Symphyotrichum anticostense, une espèce décaploïde de la région du golfe du Saint-Laurent (Québec, Nouveau Brunswick, et Maine). Il a été proposé que S. anticostense serait une allopolyploïde dérivée de l'hybride entre S. novi-belgii, une hexaploïde et S. boreale, une espèce putatifment tetraploïde. S. novi-belgii est distribuée dans la région côtière d'est d'Amérique du Nord et S. boreale est répandue à travers de la partie du nord d'Amérique du Nord de l'ouest à l'est. Ces trois espèces appartient au genre Symphyotrichum qui comprend 91 espèce. Plus que 50% de celles-ci sont polyploïdes. Les relations phylogénétiques au sein du genre sont aussi inconnues même au niveau diploïdie. Avant d'examiner l'origine de S. anticostense, il semble nécessaire dans la première étape d'expliquer les relations phylogénétiques du genre. Les objectifs de cette thèse sont donc de délimiter des espèces excluant des polyploïdes et examiner l'origine de S. anticostense. Le résultat d'analyse de donnée de l'ITS supporte presque des classifications auparavant basées sur les approches morphologiques et cytologiques. Le résultat du gène nucléaire GAPDH est faiblement en accord avec le résultat obtenu de l'ITS. Le triage des lignées incomplet dans Symphyotrichum est souligné par le triage au hasard des polymorphismes ancestraux parmi les alleles de GAPDH. Cette preuve ainsi que la résolution insuffisante parmi ribotypes, suggèrent une évolution récente du genre. Des analyses univariées et multivariées de données morphologiques ainsi que le résultat de marqueur d'ITS supportent la position taxonomique de S. anticostense comme un hybride dérivée de S. llovi-belgii et S. boreale. L'analyse basée sur réseau des ribtypes suggère trois origines géographiques pour S. anticostense: 1) Lac Saint-Jean; 2) Gaspèsie- Nouveau Brunswick- Maine; 3) l'Ile d'Anticosti. Les résultats de deux gènes nucléaires (GAPDH v et MIPS) démontrent relativement l'incongruence entre les arbres phylogenetiques. Bien que les résultats obtenus de ces marqueurs supportent ceux-là obtenus des données morphologiques et l'ITS, le triage des lignées incomplet a affecté extrêmement les relations phylogenetiques parmi des espèces. Mots clés: Asteraceae, Astereae, Symphyotrichum, Symphyotrichum Qnticostense, polyploïdie, hybridation, phylogénie, incongruence, triage des lignées incomplet, radiation. VI Table of Contents Abstract .............................................................................................................................. iii Résumé ................................................................................................................................ v Table of Contents .................................................................................................. ;........... vii List of tables ........................................................................................................................ x List of figures ..................................................................................................................... xi \ List of abbreviations ........................................................................................................ xiii Acknowledgments ............................................................................................................ xiv CHAPTER 1 Introduction ................................................................................................... 1 1.1 Taxonomic history of Symphyotrichum anticostense ................................................... 4 1.2 B iological characteristics .............................................................................................. 5 1.2.1 Taxonomy .......................................................................................................... 5 1.2.2 Distribution ...................................................................................................... 10 1.2.3 Ecology ............................................................................................................ 12 1.3 Objectives ................................................................................................................... 13 1.4 Polyploidy ................................................................................................................... 15 1.5 Hybridization .............................................................................................................. 16 1.5.l Detection ofhybrid speciation ........................................................................
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