Misumena Vatia (Thomisidae), Qui Chassent Immobiles Sur Les Fleurs Et Qui Sont Capables De Changer De Couleur

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Misumena Vatia (Thomisidae), Qui Chassent Immobiles Sur Les Fleurs Et Qui Sont Capables De Changer De Couleur UNIVERSITÉ FRANÇOIS - RABELAIS DE TOURS ÉCOLE DOCTORALE SANTE, SCIENCES, TECHNOLOGIES INSTITUT DE RECHERCHE SUR LA BIOLOGIE DE L’INSECTE THÈSE présentée par : Jérémy DEFRIZE soutenue le : 25 Juin 2010 pour obtenir le grade de : Docteur de l’université François - Rabelais Discipline/ Spécialité : Sciences de la Vie Camouflage chez les araignées crabe: Approche sensorielle, comportementale et écologique THÈSE dirigée par : M. CASAS Jérôme Professeur, Université François-Rabelais, Tours RAPPORTEURS : M. Barth Friedrich Professeur, Université de Vienne, Autriche M. Merilaita Sami Chargé de Recherche, Université d’Abo, Finland JURY : M. Barth Friedrich Professeur, Université de Vienne, Autriche M. Casas Jérôme Professeur, Université François-Rabelais, Tours M. Lazzari Claudio Professeur, Université François-Rabelais, Tours M. Merilaita Sami Chargé de Recherche, Université d’Abo, Finland M. Théry Marc Chargé de Recherche, Muséum, Paris 1 A mes parents 2 Remerciements Je tiens à remercier en premier lieu mon directeur de thèse Jérôme Casas pour m‟avoir permis de réaliser mon doctorat sur un sujet qui m‟a passionné et de m‟avoir accueilli dans l‟équipe relations multitrophiques. Je le remercie également pour son aide précieuse, ses conseils avisés et sa disponibilité sans faille durant ces quatre années. Je remercie sincèrement Claudio Lazzari pour l‟aide qu‟il m‟a apporté non seulement pendant ma thèse mais aussi pendant mon stage de Master 2 effectué dans son équipe. Je le remercie également d‟avoir accepté d‟être dans le jury de ma thèse. Je remercie Marc Théry du Muséum d‟Histoire Naturelle de Paris pour son aide et ses conseils pour le 1er chapitre de ma thèse et d‟avoir accepté d‟être membre du jury de thèse. Je remercie Eric Warrant et Almut Kelber de l‟université de Lund (Suède) pour l‟accueil chaleureux pendant mon séjour dans leur laboratoire, leurs conseils et leur aide matérielle et scientifique sur la partie électrophysiologie de ma thèse. La réalisation de cette thèse n‟aurait pu être possible aussi sans l‟aide scientifique et matérielle de nombreuses personnes de l‟IRBI : David Giron, Christelle Suppo-Magal, Sylvain Pincebourde, Térésita Insausti, Fabrice Vannier et Carolle Labrousse. Merci aussi à Clément, Sébastien, Denis, Sylvain, Chloé pour tous les bons moments passés à l‟IRBI. Je remercie l‟ensemble de l‟équipe relations multitrophiques et de l‟IRBI pour la bonne ambiance au quotidien, propice aux échanges et au travail. Je remercie Christine Besse, Helene, et Sonia Djaoui pour leur efficacité dans les taches administratives et leur gentillesse. Un grand Merci à Denis, Julien, Thomas et Germain pour les soirées ligue des Champions, ainsi qu‟à Sylvain, Aurélie, Sébastian, Romina, Arnold, Fabienne et Thomas pour l‟ensemble des soirées pendant cette thèse. Je remercie également l‟ensemble des participants assidus et accros au foot d‟Englos tous les samedi après-midi pour leur bonne humeur et les bons moments qu‟ils m‟ont fait passér toutes ces dernières années. Un grand merci à Tom H, Ju Dek, Loic, Ced, Jo Thomas, David D. dit „TP‟, Ben K. pour rendre mes (trop rares) WE à Lille si fatiguants mais aussi tellement précieux. Je n‟oublie évidemment pas les membres du carré magique Cyr‟, Tim et Chang sans qui tout serait un peu différent. 3 Enfin, je voudrais remercier chaleureusement tous les membres de ma famille: mes grands parents, mes parents, mes frères et sœurs, Mél, Caroline, Maité et Dayvi ainsi qu‟Arnaud, Thomas et mes petits neveux, Gaspard, Baptiste et Gabriel pour leur soutient à bien des niveaux. 4 RESUME De nombreuses interactions interspécifiques de type proie-prédateur font intervenir la manipulation des signaux sensoriels. Certaines espèces sont capables par exemple de produire un contraste chromatique et/ou achromatique suffisamment faible sur leur substrat pour rendre difficile leur détection par les proies ou les prédateurs, une stratégie appelée „background matching‟. Parmi ces espèces, on trouve des araignées crabe, comme Misumena vatia (Thomisidae), qui chassent immobiles sur les fleurs et qui sont capables de changer de couleur. Il est supposé depuis plus d‟un siècle que ce changement de couleur permet de devenir invisible aux yeux des proies. Mais, comme la plupart des espèces reportées comme camouflées, elles le sont de façon subjective puisque l‟appréciation du degré de camouflage est basée sur notre propre vision. Globalement, il existe un décalage entre la quantité de travaux sur l‟écologie de M. vatia, sa notoriété en tant qu‟experte du camouflage, et la connaissance réelle sur le camouflage et le changement de couleur chez cette espèce. L‟objectif de cette thèse était d‟aborder le camouflage chez M. vatia d‟un point de vue sensoriel, à une échelle communautaire, avec une approche combinant physiologie et comportement. L‟utilisation de la spectroradiométrie et des modèles de vision des couleurs ont permis de quantifier le niveau de contraste chromatique et achromatique de l‟araignée sur son substrat dans les systèmes visuels de différentes proies comme les abeilles, les bourdons, les diptères et des prédateurs comme les passériformes. Il a été ainsi démontré que si M. vatia était indétectable dans l‟achromatique à longue distance, le niveau de contraste chromatique à courte distance était fortement dépendant du substrat et de l‟identité du receveur. Des méthodes d‟électrophysiologie intra- et extracellulaire combinées à des adaptations sélectives ont permis de révéler que M. vatia possédait deux types de photorécepteurs dans chaque paire d‟yeux, l‟un ayant un pic de sensibilité dans la région des UV-A (environ 340 nm) et le second dans la région verte du spectre (environ 525 nm). Ces résultats impliquent que M. vatia possède les bases physiologiques nécessaires à la vision des couleurs, et constituent des éléments importants pour la compréhension des mécanismes visuels à l‟origine du changement de couleur. D‟un point de vue comportemental, des tests de choix floraux ont suggérés une préférence d‟ordre chromatique chez les araignées juvéniles. Cette préférence les rend peu discriminable pour les proies. Les résultats de cette thèse sont finalement replacés dans un contexte évolutif et physiologique plus général. Mots-clés: Misumena vatia, Camouflage, Ecologie sensorielle des communautés, Contraste chromatique, Contraste achromatique, Préférence colorée, Sensibilités spectrales 5 ABSTRACT Numerous prey-predator interactions rely on the manipulation of sensory signals. Some species for example are able to produce a low chromatic and/or achromatic contrast on their substrates, making them quite difficult to detect for prey or predators. This specific strategy is known as background matching. Among those species, some crab spiders, as Misumena vatia (Thomisidae), hunt motionless on flowers waiting for flower-visiting insects and are able to change its colouration from white to yellow over several days. It has assumed for more than a century that this species changes colour to decrease the risk of detection by prey and predator, such as birds. However, as for the majority of species reported as cryptic, this assumption is based only on our own subjective perception of the degree of crypsis. Overall, there is a discrepancy between the high number of works on its ecology, its fame as an expert of camouflage and the real empirical knowledge about its crypsis and colour change mechanisms. The aim of this thesis was therefore to study crypsis in M. vatia, from community sensory perspective, using an approach combining physiology and behaviour. The use of spectroradiometry and colour vision models allowed us to rigorously quantify the chromatic and chromatic contrasts of spiders in the perspective of all receivers. We showed that if M. vatia was undetectable at long distance through achromatic vision, the chromatic contrast value is quite dependent of both substrate and receiver identity. Intra- and extracellular electrophysiological recordings combined with selective adaptation have disclosed that M. vatia possesses two classes of photoreceptors in each pair of eyes: one in the ultraviolet (around 340 nm) and one in the green region (around 525 nm). M. vatia has therefore the physiological basis at the retinal level required for colour vision. Behavioural choice experiments with third instar spiderlings between artificial flowers with different chromatic and achromatic compositions suggested a chromatic preference. This preference induced a low discriminability of spiderlings on their substrate from the perspective of hymenoptera prey. In the final part, we discuss these results in an overall evolutionary and physiological context. Key-words: Misumena vatia, Crypsis, Community sensory ecology, Chromatic contrast, Achromatic contrast, Colour preference, Spectral sensitivities 6 TABLE DES MATIERES Remerciements…………………………………………………………………………......... 3 Résumé……………………………………………………………………………………..... 5 Abstract…………………………………………………………………………………........ 6 Table des matières…………………………………………………………………………… 7 Liste des tableaux……………………………………………………………………………. 10 Liste des figures……………………………………………………………………………… 11 Liste des annexes…………………………………………………………………………….. 15 INTRODUCTION GENERALE……………………………………………………………. 16 CHAPITRE 1: Camouflage chez l‟araignée crabe Misumena vatia: Une perspective d‟écologie sensorielle des communautés……………………………………………………. 25 Background colour matching by a crab spider in the field: A community sensory ecology perspective……………………………………………………………………………............ 26 Abstract…………………………………………………………………………………...
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