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Effect of Host-Plant Use on Speciation and Parasitoid Community Structure

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Effect of Host-Plant Use on Speciation and Parasitoid Community Structure dissertations Sanna Leppänen Effect of host-plant use on | Sanna| 154 Leppänen | speciation and parasitoid community structure in internal-feeding sawflies Effect use on speciation host-plant of and parasitoid community structure internal-feeding in sawflies Sanna Leppänen Effect of host-plant use on speciation and parasitoid community structure in internal-feeding sawflies Publications of the University of Eastern Finland Dissertations in Forestry and Natural Sciences Publications of the University of Eastern Finland Dissertations in Forestry and Natural Sciences isbn: 978-952-61-1560-3 (printed) issnl: 1798-5668 issn: 1798-5668 isbn 978-952-61-1561-0 (pdf) issnl: 1798-5668 issn: 1798-5676 SANNA LEPPÄNEN Effect of host-plant use on speciation and parasitoid community structure in internal-feeding sawflies Publications of the University of Eastern Finland Dissertations in Forestry and Natural Sciences 154 Academic Dissertation To be presented by permission of the Faculty of Science and Forestry for public examination in the Auditorium N100 in Natura Building at the University of Eastern Finland, Joensuu, on September, 19, 2014, at 12 o’clock. Department of Biology Author’s address: University of Eastern Finland Department of Biology P.O.Box 111 80101 JOENSUU FINLAND email: [email protected] Supervisors: Associate professor Tommi Nyman, Ph.D. University of Eastern Finland Department of Biology P.O.Box 111 80101 JOENSUU FINLAND email: [email protected] Professor Heikki Roininen, Ph.D. University of Eastern Finland Department of Biology P.O.Box 111 80101 JOENSUU FINLAND email: [email protected] Reviewers: Docent Niklas Janz, Ph.D Stockholm University Department of Zoology 106 91 STOCKHOLM SWEDEN Grano email: [email protected] Joensuu, 2014 Research scientist Carlos Lopez-Vaamonde, Ph.D Editors: Prof. Pertti Pasanen, Institut National de la Recherche Aqronomique (INRA) Prof. Pekka Kilpeläinen, Prof. Kai Peiponen, Prof. Matti Vornanen Zoologie Forestière 45075 ORLÉANS Distribution: FRANCE Eastern Finland University Library / Sales of publications email: [email protected] P.O.Box 107, FI-80101 Joensuu, Finland Opponent: Professor Niklas Wahlberg, Ph.D tel. +358-50-3058396 University of Turku http://www.uef.fi/kirjasto Department of Biology 20014 TURKU FINLAND ISBN: 978-952-61-1560-3 (printed) email: [email protected] ISSNL: 1798-5668 ISSN: 1798-5668 ISBN 978-952-61-1561-0 (PDF) ISSNL: 1798-5668 ISSN: 1798-5676 Author’s address: University of Eastern Finland Department of Biology P.O.Box 111 80101 JOENSUU FINLAND email: [email protected] Supervisors: Associate professor Tommi Nyman, Ph.D. University of Eastern Finland Department of Biology P.O.Box 111 80101 JOENSUU FINLAND email: [email protected] Professor Heikki Roininen, Ph.D. University of Eastern Finland Department of Biology P.O.Box 111 80101 JOENSUU FINLAND email: [email protected] Reviewers: Docent Niklas Janz, Ph.D Stockholm University Department of Zoology 106 91 STOCKHOLM SWEDEN Grano email: [email protected] Joensuu, 2014 Research scientist Carlos Lopez-Vaamonde, Ph.D Editors: Prof. Pertti Pasanen, Institut National de la Recherche Aqronomique (INRA) Prof. Pekka Kilpeläinen, Prof. Kai Peiponen, Prof. Matti Vornanen Zoologie Forestière 45075 ORLÉANS Distribution: FRANCE Eastern Finland University Library / Sales of publications email: [email protected] P.O.Box 107, FI-80101 Joensuu, Finland Opponent: Professor Niklas Wahlberg, Ph.D tel. +358-50-3058396 University of Turku http://www.uef.fi/kirjasto Department of Biology 20014 TURKU FINLAND ISBN: 978-952-61-1560-3 (printed) email: [email protected] ISSNL: 1798-5668 ISSN: 1798-5668 ISBN 978-952-61-1561-0 (PDF) ISSNL: 1798-5668 ISSN: 1798-5676 ABSTRACT differentiation has a huge impact on galler speciation, but also that host-associated differentiation is not completely repeatable: Plants, herbivorous insects, and parasitoids form a major part of the two galler genera have similar host plant repertoires, but ΦST all described species. Still, the evolutionary processes that have estimates among population samples collected from specific led to the formation of these diverse food webs with extremely willow species differ markedly. complex multitrophic interactions are not fully understood. Parasitoids of both leafminers and gallers represent three Insect–plant interactions are taxonomically conserved in the hymenopteran families that are phylogenetically distant from sense that herbivores tend to use closely related host plants, but each other. In the case of miner parasitoids, closely related host–plant shifts are common in most insect taxa. There is parasitoids tend to use the same miner species. However, evidence that shifts among host plants can influence insect phylogeny-based analyses show that the host-plant phylogeny speciation, but the effects of niche shifts on higher trophic levels is a more important factor structuring parasitoid–miner remain a mystery. In addition, the third trophic level (especially associations than is the miner phylogeny. The diet breadth of parasitoids) could be the driving force behind insect individual parasitoid species varies, and both niche- and diversification by causing insects to shift host plants. Using species-specialist enemies can be found. In combination, the molecular-genetic methods, this thesis tries to answer how host tritrophic patterns suggest that both bottom–up and top–down plants influence speciation in leaf-mining and gall-inducing forces have influenced diversification in the plant–leafminer– sawflies and in their associated parasitoids. My main purpose parasitoid system. was to investigate the determinants of tritrophic associations, DNA barcoding proved to be a suitable tool for studying the and the role of parasitoids in herbivore speciation. structure of leaf-galler parasitoid communities. In galler A molecular-phylogenetic analysis of leaf-mining sawflies in parasitoids, habitat (boreal–subarctic vs. arctic–alpine) had the the subfamily Heterarthrinae indicates that the subfamily strongest effect on parasitoid community structure. This originated c. 100 million years ago, which means that the indicates that, in addition to host shifts, colonization of novel leafminers are younger than their main host taxa. habitats may provide shelter from enemies and accelerate Reconstructions of larval feeding modes show that transitions herbivore speciation. Furthermore, at least two parasitoid from external to internal feeding have occurred multiple times species apparently originated by tracking galler hosts that had within the superfamily Tenthredinoidea, but also that leaf- shifted to using dwarf willows that grow in treeless arctic– mining has arisen only twice. On long time scales, host–plant alpine habitats. use is taxonomically unstable, and multiple convergent host In combination, the results of the leafminer and galler studies shifts have happened during the evolutionary history of support the view that parasitoids could be a major driving force Heterarthrinae. in herbivore speciation, but also that diversification can cascade Host-plant shifts also seem to promote genetic differentiation up in the food web, when parasitoids form host races attacking and speciation in gall-inducing sawflies belonging to the genera herbivores on novel plant species and niches. Pontania and Euura, as clear host-based clustering of individuals is observable in phylogenetic trees. Indeed, hierarchical Universal Decimal Classification: 575.858, 575.86, 591.522, 591.531.1, AMOVAs indicated that most of the genetic variation in both 591.557.8, 595.793 galler genera is explained by willow host species rather than by geographic location. The results show that host-associated ABSTRACT differentiation has a huge impact on galler speciation, but also that host-associated differentiation is not completely repeatable: Plants, herbivorous insects, and parasitoids form a major part of the two galler genera have similar host plant repertoires, but ΦST all described species. Still, the evolutionary processes that have estimates among population samples collected from specific led to the formation of these diverse food webs with extremely willow species differ markedly. complex multitrophic interactions are not fully understood. Parasitoids of both leafminers and gallers represent three Insect–plant interactions are taxonomically conserved in the hymenopteran families that are phylogenetically distant from sense that herbivores tend to use closely related host plants, but each other. In the case of miner parasitoids, closely related host–plant shifts are common in most insect taxa. There is parasitoids tend to use the same miner species. However, evidence that shifts among host plants can influence insect phylogeny-based analyses show that the host-plant phylogeny speciation, but the effects of niche shifts on higher trophic levels is a more important factor structuring parasitoid–miner remain a mystery. In addition, the third trophic level (especially associations than is the miner phylogeny. The diet breadth of parasitoids) could be the driving force behind insect individual parasitoid species varies, and both niche- and diversification by causing insects to shift host plants. Using species-specialist enemies can be found. In combination, the molecular-genetic methods, this thesis tries to answer how host tritrophic patterns suggest that both bottom–up and top–down plants influence speciation in leaf-mining
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