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Thèse Comlpète Pour 14 Décembre MINISTÈRE DE L’ALIMENTATION DE L’AGRICULTURE ET DE LA PÊCHE MONTPELLIER SUPAGRO THÈSE présentée à Montpellier SupAgro pour obtenir le diplôme de Doctorat Formation doctorale : Évolution, Écologie, Ressources génétiques, Paléontologie École doctorale : Systèmes Intégrés en Biologie, Agronomie, Géosciences, Hydrosciences, Environnement Laboratoire d’accueil : Unité Mixte de Recherche Centre de Biologie pour la Gestion des Populations INRA / IRD / CIRAD / Montpellier SupAgro Campus International de Baillarguet, CS 30016 34988 Montferrier-sur-Lez cedex, France Etudes taxonomiques de deux genres d’acariens prédateurs de la famille des Phytoseiidae (Acari : Mesostigmata) : Phytoseiulus Evans et Neoseiulella Muma Présentée et soutenue publiquement par MOHAMAD KANOUH le 14 Décembre 2010 JURY Serge KREITER Montpellier SupAgro Co-directeur de thèse Mark JUDSON Muséum National d’Histoire Naturelle, Paris Examinateur Jean-Pierre LUMARET Université de Montpellier III Examinateur André NEL Muséum National d’Histoire Naturelle, Paris Rapporteur Jean-Loup NOTTEGHEM Montpellier SupAgro Examinateur Salvatore RAGUSA Université degli Studi de Palermo, Italie Rapporteur Marie-Stéphane TIXIER Montpellier SupAgro Co-directrice de thèse RÉSUMÉ La classification actuelle de la famille des Phytoseiidae n’est pas basée sur de réelles études phylogénétiques et par conséquent, de nombreuses questions se posent sur la validité des taxa supra-spécifiques mais également vis-à-vis des taxa spécifiques. Ce travail de thèse avait donc pour objectif de répondre à de telles questions pour deux genres : Phytoseiulus et Neoseiulella, en utilisant pour la première fois des approches phylogénétiques moléculaires et morphologiques. Ces études phylogénétiques ont été également associées à des études biogéographiques. Les résultats obtenus par ces deux approches sont congruents et semblent montrer que ces deux genres ne sont pas monophylétiques : le genre Phytoseiulus semble au mieux paraphylétique, tandis que le genre Neoseiulella serait polyphylétique. Ces résultats sont différents de ceux développés dans les diverses révisions successives de ces deux taxa. Ce travail de thèse a permis de répondre également à des questionnements sur cinq synonymies au sein du genre Neoseiulella . L’étude de pratiquement tous les types des espèces du genre Neoseiulella nous a permis de redéfinir le genre, en excluant trois espèces et en discutant certaines synonymies. Une clé d'identification des espèces valides du genre Neoseiulella basée sur les adultes femelles a également été proposée. Des études complémentaires, notamment des analyses moléculaires, seront nécessaires afin de conclure plus définitivement sur les relations évolutives entre les taxa étudiés. MOTS CLEF : Phytoseiidae, Phytoseiulus , Neoseiulella , phylogénie, biogéographie ABSTRACT The systematic of the family Phytoseiidae is not presently based on solid phylogenetic studies and therefore, many taxonomic questions still arise, concerning the validity of supra-specific and specific taxa identified to-date. This thesis thus aimed to answer such questions for two genera, Phytoseiulus and Neoseiulella , using for the first time molecular and morphological phylogenetic analyses. Biogeographic analyses have been also carried out. Results obtained by both morphological and molecular approaches are congruent and seem to show that both genera are not monophyletic: Phytoseiulus seems paraphyletic whereas Neoseiulella seems polyphyletic. These results are different from those encountered in previous revisions of these two taxa. Furthermore, this study allowed to conclude on five synonymies within the genus Neoseiulella . The observation of nearly the totality of the type materials of the genus Neoseiulella permitted to redefine this genus, excluding three species and discussing some synonymies. Lastly, an identification key to the valid species of the genus Neoseiulella based on adults females was proposed. Further experiments, including molecular investigations, are however still required in order to obtain more reliable conclusions on the evolutionary relationships of the studied taxa. KEY WORDS: Phytoseiidae, Phytoseiulus , Neoseiulella , phylogeny, biogeography I REMERCIEMENTS Un grand MERCI à tous celles et ceux à qui ont partagé avec moi leurs connaissances, qui ont investi de leur temps et de leur énergie pour que ce projet aboutisse: Je voudrais tout d'abord exprimer mes remerciements à mes encadrants de thèse : Madame Marie-Stéphane Tixier Lopes et Monsieur Serge Kreiter , pour m'avoir accueilli dans leur laboratoire, pour avoir dirigé mon travail depuis le Master II; pour leur soutien constant tout au long de ces études, pour leur gentillesse et leur confiance pendant ces années de travail. Je souhaite exprimer toute ma gratitude envers les membres du jury, M. André NEL, M Mark JUDSON, M. Jean-Loup NOTTEGHEM, M. Jean-Pierre LUMARET, et M. Salvatore RAGUSA d’avoir accepter de juger ce travail présent. Mes remerciements à mon gouvernement syrien de m’avoir donné la chance d’être parmi vous ce jour là, et d’avoir financé ce travail de thèse, et également à ceux qui m’ont facilité ce long chemin, M. Majd Jamal et M. Mohammed Adel Ftaeyh. Je souhaite exprimer mes remerciements à Gaël Kergoat et Jean François Martin pour leurs aides et leurs conseils précieux dans la démarche phylogénétique. Merci à Brigitte Cheval , Sabine Guichou , Philippe Auger et Martial Douin pour leurs aides, leur sympathie et leur amitié. Merci également à Mireille Okassa et Maxime Ferrero de m’avoir accompagné durant ces années de thèse, pour leur amitié . Merci à toutes les personnes à SupAgro et au CBGP avec qui j’ai partagé de bons moments. Je dédie ce travail à mes parents, sans qui je ne serais pas où j’en suis aujourd’hui ; Merci à tout membre de ma famille, qui m’ont toujours soutenu. Finalement, je ne trouve pas les mots qu’il faut pour exprimer mes remerciements du fond du mon cœur à mon épouse Feryal et à mes deux "anges" Hamza et Ammar … MERCI ! II PUBLICATIONS INCLUSES DANS LA THÈSE A1 – KANOUH M, TIXIER M-S, OKASSA M, KREITER S, 2010. Phylogenetic and Biogeographic analysis of the genus Phytoseiulus (Acari : Phytoseiidae). Zoologica Scripta, 39 (5), 450-461. A2 – KANOUH M, TIXIER M-S, GUICHOU S, CHEVAL B, KREITER S, 2010. Two synonymy cases within the genus Neoseiulella (Acari : Phytoseiidae): is the molecular evidence so evident? Biological Journal of Linean Society , 101, 323-344. A3 – KANOUH M, KREITER S, TIXIER M-S, 2010. Revision of the genus Neoseiulella Muma (Acari : Phytoseiidae) with synonymies assessment and additional morphological data. A soumettre à International Journal of Acarology avant janvier 2011. LISTE DES ANNEXES Annexe 1 - Répartition géographique détaillée des espèces du genre Phytoseiulus . Annexe 2 - Les 47 espèces du genre Neoseiulella figurant dans les révisions successives. Annexe 3 - Répartition géographique détaillée des espèces du genre Neoseiulella . Annexe 4 - Les différentes familles botaniques des espèces de plantes supports sur lesquelles les espèces du genre Neoseiulella ont été signalées. Annexe 5 - Moyennes des 32 caractères morphologiques continus des espèces du genre Neoseiulella utilisés pour réaliser l’Analyse en Composantes Principales (ACP). Annexe 6 - Etat de caractères des 25 caractères morphologiques discrets des espèces du genre Neoseiulella utilisés pour réaliser l’Analyse Factorielle des Correspondances (AFC). Annexe 7 - Les 12 groupes d’espèces constitués au sein du genre Neoseiulella , représentant les différents états de 21 caractères morphologiques discrets utilisés pour réaliser des analyses phylogénétiques au sein de la sous-famille des Typhlodrominae. III LISTE DES FIGURES Figure 1. Position systématique de la famille des Phytoseiidae (d’après Lindquist et al., 2009) ................................................................................................................................. 5 Figure 2. Morphologie et dentition des chélicères des adultes (a) femelle et (b) mâle des Phytoseiidae............................................................................................................. 6 Figure 3. Face dorsale théorique d’une femelle de Phytoseiidae montrant la chaetotaxie dorsale et les différentes paires de solénostomes (gd)............................................. 8 Figure 4. Face ventrale des adultes (a) femelle et (b) mâle de Phytoseiidae........................... 10 Figure 5. Caractéristiques de la spermathèque des femelles de Phytoseiidae : (a), détails de la spermathèque; (b), différentes formes de spermathèques chez les Phytoseiidae .. 11 Figure 6. Cycle de développement des Phytoseiidae à 27 °C et 70 % d'humidité relative (d’après Yaninek et al ., 1989) ............................................................................... 12 Figure 7. Détails des caractères morphologiques d’une femelle adulte type (a-d) du genre Phytoseiulus : a, face dorsale; b, face ventrale; c, chélicère; d, spermathèque; et le spermathodactyle d’un mâle adulte (e).................................................................. 21 Figure 8. Détails des caractères morphologiques d’une femelle de Afroseiulus robertsi (Baker) : a, face dorsale; b, face ventrale; c, spermathèque; d, chélicère; e, patte IV........................................................................................................................... 23 Figure 9. Femelle de Phytoseiulus persimilis attaquant (a) une femelle et (b) un œuf de Tetranychus urticae ..............................................................................................
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