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Community-Level Effects of Non-Indigenous Aquatic Ecosystem

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Community-Level Effects of Non-Indigenous Aquatic Ecosystem Community-level effects of nonindigenous aquatic ecosystem engineers Jessica MacKay Ward Department of Biology McGill University Montreal, Canada January 2010 A thesis submitted to McGill University in partial fulfillment of the requirements of the degree of Doctor of Philosophy © Jessica MacKay Ward 2010 Library and Archives Bibliothèque et Canada Archives Canada Published Heritage Direction du Branch Patrimoine de l’édition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 Ottawa ON K1A 0N4 Canada Canada Your file Votre référence ISBN: 978-0-494-68563-1 Our file Notre référence ISBN: 978-0-494-68563-1 NOTICE: AVIS: The author has granted a non- L’auteur a accordé une licence non exclusive exclusive license allowing Library and permettant à la Bibliothèque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par télécommunication ou par l’Internet, prêter, telecommunication or on the Internet, distribuer et vendre des thèses partout dans le loan, distribute and sell theses monde, à des fins commerciales ou autres, sur worldwide, for commercial or non- support microforme, papier, électronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L’auteur conserve la propriété du droit d’auteur ownership and moral rights in this et des droits moraux qui protège cette thèse. Ni thesis. Neither the thesis nor la thèse ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent être imprimés ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author’s permission. In compliance with the Canadian Conformément à la loi canadienne sur la Privacy Act some supporting forms protection de la vie privée, quelques may have been removed from this formulaires secondaires ont été enlevés de thesis. cette thèse. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n’y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. Abstract The identification of generalities regarding which biological invasions are most likely to cause major ecological disruptions is a central goal of invasion ecology, having both theoretical and applied importance. Introduced ecosystem engineers (organisms that modify, create and/or destroy habitats) can exert strong impacts on recipient communities, altering not only species composition but also the physical structure of ecosystems. In this thesis, I examine variation in the impacts of nonindigenous ecosystem engineers on aquatic macroinvertebrate communities, with emphasis given to Dreissena spp. because they are well- studied exotic ecosystem engineers of both ecological and economic concern. The objectives of this research were to identify general patterns and test existing theories with respect to the community-level impacts of exotic ecosystem engineers. To meet these objectives, I employed a combination of field experiments and statistical syntheses of published data. In Chapters 2 and 3, I examined the reliance of ecosystem engineering impacts on a community‘s evolutionary experience with the engineer or with similar species (i.e. ecological naiveté). Meta-analyses revealed that impacts on community diversity and abundance do not systematically differ among native and exotic ecosystem engineers (Chapter 2), and suggested that introduced ecosystem engineers with distinct functional traits compared to native species generally promote facilitation (Chapter 3). I focused on Dreissena spp. in Chapters 4 and 5, wherein I examined sources of variation in the community-level impacts of these exotic mussels. A meta-analysis revealed that Dreissena generally increases macroinvertebrate density and richness, and that the strength of these effects varies with sediment particle size (Chapter 4). In Chapter 5, I tested the effect of Dreissena patch topography on benthic diversity by experimentally manipulating the area and perimeter-to-area ratio of mussel patches. My results indicated that Dreissena patch topography may be an important determinant of species richness at small spatial scales, and suggested that the impact of Dreissena is modified in the presence of another habitat-forming species, the native alga Cladophora. This i work contributes to our understanding of the context-dependence of invasion impacts; that is, how the direction or magnitude of impact depends on the characteristics of invaders and invaded systems. ii Résumé L'identification de généralités en ce qui concerne les invasions biologiques d‘qui ont les plus grandes chances de provoquer des perturbations écologiques importantes est un but central de l'écologie des invasions, ayant de l'importance théorique et pratique. Les ingénieurs d'écosystèmes introduits (les organismes qui modifient, créent et-ou détruisent des habitats) peuvent exercer des impacts importants sur les communautés envahies, en transformant non seulement la composition d'espèces, mais aussi la structure physique des écosystèmes. Dans cette thèse, j'examine la variation dans les impacts d'ingénieurs d'écosystème nonindigènes sur les communautés des macroinvertébrés aquatiques, avec l'accentuation donnée à Dreissena spp. parce qu'elles sont bien étudiées en tant qu'ingénieurs d'écosystèmes qui ont des impacts écologiques et économiques importants. Les objectifs de cette recherche sont d'identifier des généralités et d‘évaluer des théories en ce qui concerne les impacts d‘ingénieurs d'écosystèmes exotiques au niveau de la communauté. Pour atteindre ces objectifs, j'ai employé une combinaison d‘expériences sur le terrain et de synthèses statistiques de données publiées. Dans les chapitres 2 et 3, j'ai examiné la dépendance des impacts d'ingénieurs d‘écosystèmes sur l'expérience évolutive d'une communauté soit avec la présence de l'ingénieur ou bien avec des espèces semblables (c'est-à- dire la naïveté écologique). Les meta-analyses ont démontré que les impacts sur la diversité et l'abondance communautaires ne diffèrent pas systématiquement parmi les ingénieurs d'écosystème indigènes et exotiques (chapitre 2), et ont suggéré que les ingénieurs d'écosystème exotiques qui possèdent des traits fonctionnels distinctifs comparés aux espèces indigènes favorisent généralement la facilitation (chapitre 3). Je me suis concentrée sur les Dreissena spp. dans les chapitres 4 et 5, où j'ai examiné les sources de variation dans les impacts de ces moules exotiques au niveau communautaire. Une meta-analyse a démontré que Dreissena augmente généralement la densité et la richesse des macroinvertébrés, et que l‘ampleur de ces effets varie selon la granulométrie des particules de sédiment (chapitre 4). Dans le chapitre 5, j'ai évalué l'effet de la topographie de iii parcelles de Dreissena sur la biodiversité des macroinvertébrés benthiques en manipulant expérimentalement la surface occupée et le rapport du périmètre et de la surface des parcelles de moules. Mes résultats ont indiqué que la topographie des parcelles de Dreissena est peut-être un déterminant important de la richesse des espèces aux échelles spatiales petites et ont suggéré que l'impact de Dreissena est modifié en présence d'une autre espèce créateur d‘habitats, l'algue indigène Cladophora. Ce travail contribue à notre compréhension de la dépendance vis-à- vis du contexte des impacts d'invasions; c'est-à-dire la manière dont la direction ou l‘ampleur d'impact dépend des caractéristiques des envahisseurs et des systèmes envahis. iv Acknowledgements I am above all grateful to my supervisor, Dr. Anthony Ricciardi, for his frequent council and creative input, enthusiasm for research, lively and stimulating discussions, and support throughout my development from an undergraduate student to a young scientist. I am also grateful to the members of my supervisory committee, Dr. Frédéric Guichard and Dr. Brian Leung, for their constructive input, questions and advice. I would also like to thank Dr. Andrew Gonzalez for constructive input and suggestions, Dr. Andrew Hendry for thoughtful questions, and Dr. Brian McGill for advice on statistical analyses. I am sincerely grateful to Michelle Palmer, who provided invaluable training and assistance with field research. I also thank Lisa Jones, Anneli Jokela, Rebekah Kipp, Åsa Kestrup, Robin Tiller, James Snider and Taïca Replansky for their assistance in the field and help processing experimental substrates in the lab. For assistance with invertebrate identification, I thank Rebekah Kipp, Arthis Sowmithran, Niloufar Bayani, and Hanie Seo. For assistance with data entry, I thank Suzanne Ellwood, Katie O‘Sullivan and Ahdia Hassan. I am also grateful to past and current members of the Ricciardi lab for stimulating discussions, creating a friendly work environment, and providing humour and support. I am genuinely grateful to my family and friends, especially James Snider, Barbara Ward, and Patricia MacKay for their continuous support and encouragement. I dedicate this thesis to them. In conducting this work, I was supported financially by a Canada Graduate Scholarships from the National Research and Engineering Council of Canada, a Vineberg Family Fellowship from the Department of Biology, and a Dr. and Mrs. Milton Leong Graduate Student Award from McGill‘s Graduate and Postdoctoral Studies Office. Travel
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