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Analyse De L'origine Des Fortes Variations D'impact De Tischeria

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Analyse De L'origine Des Fortes Variations D'impact De Tischeria article Analyse de l’origine des fortes variations d’impact de Tischeria ekebladella, micro-lépidoptère minant les feuilles de Chêne, commun en Bourgogne Jean BÉGUINOT 1,2 Résumé Comme partout ailleurs, nos Chênes bourguignons hébergent de nombreux insectes dont le stade larvaire mine l’intérieur des feuilles. L’impact de ces insectes, tel qu’il peut aisément s’apprécier par la densité des mines foliaires, est éminemment variable selon les espèces mais également d’une station à l’autre (d’un chêne à un autre) pour une même espèce mineuse. Des variations d’impact d’un ordre de grandeur et parfois bien davantage sont courantes et se rencontrent au demeurant chez la plupart des insectes phytophages. Cependant, ces variations d’impact n’ont pas ordinairement pu recevoir d’explications exhaustives, intégrant l’ensemble des contributions des facteurs susceptibles d’infl uer sur le niveau d’impact. Cette limitation ne tient pas à une insuffi sance d’approche théorique mais plutôt à la quasi-impossibilité d’évaluer en direct, sur le terrain, l’ensemble des principaux facteurs infl uents, en particulier s’agissant d’insectes de petites taille, infra-centimétrique. Le recours à une procédure permettant de restituer indirectement et rétrospectivement les valeurs estimées des facteurs gouvernant le niveau d’impact (procédure ‘MELBA’), à partir de données de terrain aisément accessibles a posteriori, permet de lever commodément cette diffi culté, au moins pour le cas particulier des insectes phytophages à stade larvaire se développant aux dépens d’une seule et même feuille hôte : insectes inducteurs de galles ou fonceurs de mines. Le micro-lépidoptère Tischeria ekebladella est l’une des espèces mineuses les plus spectaculaires sur nos chênes, sujette elle aussi à forte variation de niveau d’impact d’une station à une autre. On met ici à profi t la procédure susmentionnée pour préciser quantitativement les contributions respectives des différents facteurs infl uant sur la variabilité observée du niveau d’impact de Tischeria ekebladella. On montre notamment que les variations de qualité foliaire des chênes hôtes contribue tout autant (ici pour 49 %) aux variations d’impact de l’insecte que la densité locale des mères pondeuses (45 %), souvent seule invoquée. Un autre facteur, ordinairement totalement ignoré, le niveau de fécondité par ponte (taille de ponte élémentaire) joue également ici, avec une contribution il est vrai comparativement minoritaire (6 %). L’application de la procédure ‘MELBA’ permet en outre d’accéder indirectement à des traits de comportement inobservables en pratique. On montre ainsi que les mères sont assez exigeantes quant au niveau de qualité des feuilles de chênes acceptables pour ponte (de 50 % jusqu’à plus de 80 % des feuilles du chêne-hôte peuvent être rejetées selon les stations). En revanche, on met en évidence que la qualité foliaire n’apparaît pas avoir d’infl uence sur la taille de ponte élémentaire et n’exerce pas non plus de rôle attractif différentiel à distance sur les mères pondeuses. Mots-clés : Lepidoptera, chêne, mine, sélection, hôte. Analysing the origins of the strong variability of impact by Tischeria ekebladella, a common mining moth on oak leaves Abstract Tischeria ekebladella (Lepidoptera : Gracillariidae) is a common moth with its caterpillar often mining oak leaves, more rarely chestnut leaves. As a rule, the mining impact of the species exhibits strong variations according to locations, inviting to investigate quantitatively the respective contributions of the three main factors that regulate the level of mines density within host. As the direct monitoring in the fi eld of these governing factors is very diffi cult (when not virtually impossible), we use a newly designed procedure (“MELBA”) which allows tracing back to these factors indirectly, on the basis of posterior, easily recorded data (in terms of the histogram of the distribution of the numbers of mines per host- leaf). The respective contributions to the mine impact variability of Tischeria ekebladella, according to locations, were estimated by this procedure as: (i) 49% due to variations of host-leaf quality ‘α’, (ii) 45% due to variations of mothers’ local density (density of mothers’ visits to host-leaves ‘µ’) and (iii) a small contribution only (6%) due to variation of clutch-size ‘nc’. The estimated degree of mothers’ selectivity among host-leaves prior to egg-laying is rather strong (proportion of rejected host-leaves from 50% up to more than 80%). Yet, the host-leaf quality proves having no signifi cant infl uence upon the clutch-size in this species. Also, the average quality of leaves within a given oak tree seems to have no appeal on females and thus, no signifi cant infl uence on the local density of Tischeria mothers. Key words : Lepidoptera, oak, mine, selection, host. 1 Société d’histoire naturelle du Creusot - 12 rue des Pyrénées - 71200 Le Creusot - [email protected] 2 Université de Bourgogne, département Biogéosciences - 6 boulevard Gabriel - 21000 Dijon Revue scientifi que Bourgogne-Nature - 21/22-2015, 91-98 91 ÉGUINOT Jean B Trois mines de Tischeria ekebladella sur une feuille de chêne. IMBER Ian K , Hungary Forest Research Institute, Bugwood.org Hungary Research Institute, , Forest KA ó S György C Larve de Tischeria ekebladella. Imago de Tischeria ekebladella. 92 Jean BÉGUINOT Revue scientifi que Bourgogne-Nature - 21/22-2015, 91-98 Introduction et position du problème Parmi les choix alimentaires variés des insectes, les plantes (toutes parties comprises) mais aussi les autres insectes (tous stades compris mais surtout larvaires) sont les cibles les plus fréquemment adoptées et font donc principalement les frais de la voracité entomologique. En raison même de cette voracité, ni les plantes ni les insectes-proies ne sont restés inertes face au danger. Chacun a donc développé sa (ou ses) propre(s) tactique(s) défensive(s), au fi l de l’évolution. Ainsi, les plantes ont surtout mis au point des défenses chimiques dissuasives (alcaloïdes, tanins, etc…) tandis que les insectes- proies ont surtout développé des comportements d’évitement, incluant d’ailleurs à l’occasion la dissuasion chimique et ce, d’ailleurs en tirant souvent parti des produits toxiques d’origine végétale. Par contrecoup, les insectes consommateurs ont dû, à leur tour, s’adapter à ces moyens de défense, plus ou moins différents selon les espèces consommables, en se spécialisant très étroitement pour mieux tenter de contourner les défenses propres à la catégorie de proie fi nalement retenue au menu. Les insectes herbivores se spécialisent donc et la plupart d’entre eux sélectionnent exclusivement à l’échelle de la famille, souvent même à l’échelle du genre, voire de l’espèce de plante fi nalement acceptée au menu et se concentrent en général sur une partie délimitée de la plante. De même les insectes prédateurs (en général parasites ou parasitoïdes) se spécialisent souvent sur des catégories taxonomiques étroites de proies, se concentrant en outre sur un stade bien défi ni du développement des dites proies. Tout ceci explique la diversité spécifi que extraordinaire des insectes, conséquence directe de cette habituelle hyperspécialisation alimentaire. Encore faut il ajouter qu’à chacun de ces centaines de milliers de menus distincts peut correspondre plusieurs approches consommatrices différentes… Par exemple, considérant les herbivores, il peut s’agir simplement de croquer la plante de l’extérieur, autant que nécessaire à l’appétit. Mais des comportements plus exigeants et plus complexes se rencontrent souvent chez les petites espèces d’insectes herbivores, de taille typiquement infra-centimétrique, comme telles plus exposées aux aléas environnementaux. L’insecte demande alors à la plante hôte non seulement le couvert mais encore un gîte sûr. Il en est ainsi, typiquement, des insectes minant les feuilles ou des insectes induisant des galles par manipulation de l’expression du patri- moine génétique de la plante hôte. Pour eux, l’ensemble du développement s’effectuera le plus souvent aux dépens de la seule feuille qui les porte, celle où la mère a déposé l’œuf dont ils sont issus. Or, pour ces insectes très délicats, la qualité des feuilles d’une même plante-hôte est ressentie comme éminemment variable, lors même que ces feuilles sont pour nous absolument identiques entre-elles. À charge donc pour la mère, de réaliser successivement une série de bons tests et vérifi cations : 1) choisir la bonne espèce-hôte, puis 2) choisir la bonne partie de la plante (feuille ou tige ou racine ou fl eur ou fruit ou bourgeon de feuille ou bourgeon de fl eur…), enfi n 3) pour la partie de plante sélectionnée, tester le niveau de ‘qualité’, ne retenant fi nalement pour la ponte que les supports qui pré- sentent un niveau qualitatif suffi sant au point de vue de la mère. Rude et lourde tâche qui explique les nombreux contrôles que la mère effectue méthodiquement avant ponte. Chez ces menues espèces, la fragilité du stade larvaire demande, en compensation, des comportements souvent bien plus sophistiqués (au moins à cet égard) que chez les espèces moins ténues. Ce long préambule pour expliquer que chez les insectes mineurs de feuilles notam- ment, une part seulement des feuilles de la plante hôte en cours de visite saura satisfaire la mère pondeuse ; part que nous désignerons dans la suite par ‘α’, et qui représente donc le ratio d’acceptabilité des feuilles-hôtes par les mères pondeuses. Voyons dès lors de quoi dépend « l’impact » d’un insecte mineur donné sur une plante hôte, impact exprimé par la densité de mines, c’est-à-dire le ratio I du nombre de mines rencontrées rapporté au nombre de feuilles examinées. Pour qu’une feuille héberge une voire plusieurs mines il faut, bien entendu, qu’elle ait reçu au moins une visite exploratoire par une mère pondeuse et qu’à la suite de cette (ces) visite(s) elle ait été acceptée (probabilité α).
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