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Diversity of Canopy Spiders in North-Temperate Hardwood Forests

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Diversity of Canopy Spiders in North-Temperate Hardwood Forests NOTE TO USERS This reproduction is the best copy available. UMI* Diversity of canopy spiders in north-temperate hardwood forests Maxim Larrivee Department of Natural Resource Sciences McGill University Montreal, Quebec, Canada April 2009 A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Doctor of Philosophy © M. Larrivee 2009 Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 Ottawa ON K1A 0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-66615-9 Our file Notre reference ISBN: 978-0-494-66615-9 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque 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 telecommunication ou par I'lnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. 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. 1+1 Canada Abstract The objective of this thesis was to understand the spatial patterns and processes responsible for canopy and understorey spider (Arachnida: Araneae) diversity at multiple spatial scales in north-temperate hardwood forests. I sampled tree trunks (sticky traps) and foliage (beating) of sugar maple and American beech tree canopies and their understorey saplings in old growth forests near Montreal, Quebec. Results show the composition of canopy and understorey assemblages differed significantly, and so did sugar maple and American beech canopy assemblages. Each stratum was also dominated by different species. The rank-abundance distribution of species from each habitat wsa also verticaly stratified because it fit different distribution models. Different factors likely structure assemblages in both habitats, particularly since the canopy is a less stable environment. Spiders from canopy and understorey foliage were tested in a laboratory for their propensity to balloon. General linear models indicated that small sized web-building spiders of the RTA and Orbicularia clades have the highest propensity to balloon. Small bodied species initiated ballooning regardless of the habitat they were collected in or their developmental stage. My results support the mixed evolutionary stable strategy theory and indicate the absence of risk-spreading in the dispersal strategy of canopy spiders. My last chapter focused on dispersal capacity and diversity patterns of spiders at multiple spatial scales. Analyses of the species diversity of limited and high dispersal capacity species subsets through nested-multivariate ANOVA, additive diversity partitioning, and species-abundance distribution curves all point towards species-sorting processes as the main driver of local community spider diversity at the tree and stand spatial scales. Mass- effects and patch-dynamic processes drive site and regional scale diversity patterns. This I thesis demonstrates that spiders provide good models to test many biological hypotheses. The research chapters of this thesis test hypotheses on the vertical stratification of forest spider diversity, the evolution of local dispersal adaptations, and the importance of dispersal capacity on species diversity patterns through a metacommunity framework. II Resume L'objectif de cette these etait de comprendre les patrons spatiaux de diversite des araignees (Arachnida: Araneae) et les processus les creant a echelles spatiales multiples en forets decidues temperees nordiques. J'ai echantillonne les troncs d'arbres (pieges collants) et le feuillage (battage) dans la canopee d'erable a sucre et d'hetre a grandes feuilles ainsi que dans la haute regeneration de chaque espece dans des forets surannees dans la region de Montreal, Quebec. Les resultats indiquent que la composition des assemblages de la canopee et de la strate arbustive different significativement de meme que ceux des canopees d'erable a sucre et de hetre a grandes feuilles. Des especes differentes sont significativement plus abondantes dans chaque strate d'elevation. La distribution de rangs d'abondance des especes indique que des mecanismes differents regissent la structure des assemblages de chaque habitat. La propension au ballonnement des araignees du feuillage de la canopee et de la haute regeneration a ensuite ete testee en laboratoire. Des modeles lineaires generaux indiquent que les petites araignees tisseuses de toile des clades RTA et Orbicularia ont une forte propension au ballonnement et que cette derniere est peu affectee par la stabilite de l'habitat. Les araignees de petite taille ballonnent peu importe leur niveau de developpement. Mes resultats supportent la theorie de la strategic evolutive mixte stable et l'absence d'une repartition du risque dans la strategie de dispersion des araignees de la canopee. Mon dernier chapitre traite de P influence de la capacite de dispersion sur les patrons de diversite des araignees a echelles spatiales multiples. Des analyses multi variees et nidifiees de sous-ensembles d'especes d'araignees a capacite de dispersion reduite ou forte indiquent que des processus de tri des especes produisent les patrons de diversite des araignees aux echelles III spatiales de l'arbre et du peuplement alors que des processus d'effets de masse et des dynamiques de peuplement regissent les patrons de diversite aux echelles du site et de la region. Cette these demontre que les araignees sont excellentes pour tester plusieurs hypotheses ecologiques liees a la distribution de la diversite verticale et horizontale, et ce sur plusieurs echelles spatiales. IV Contribution of authors This thesis consists of 3 original papers co-authored by my supervisor, Dr. Christopher Buddie. Chapter 3 of this thesis is accepted for publication in the journal Agricultural and Forest Entomology. I intend to submit Chapter 4 to the journal Ecological Entomology as an original paper, and the original paper version of Chapter 5 is in preparation for the Journal of Animal Ecology. For each of the papers presented within, I planned the experimental designs with valuable input from my supervisor and my committee members Drs. A.Gonzalez and T. Wheeler. I was responsible for fieldwork, identification, and I performed all analyses presented with feedback from my supervisor and committee members. I wrote the first drafts of all manuscripts. Final drafts contained useful comments and at times direct contributions of my supervisor. All others who contributed to the production of these papers through field or laboratory work, or through an input of ideas or financial support are acknowledged within each original paper. V Contributions to knowledge Chapter 3: This original paper characterized and compared foliage spider diversity in the canopy and understorey of sugar maple and American beech in old-growth north-temperate hardwood forests. Several aspects of this original paper are novel contributions to canopy ecology and spider ecology: 1) Use of an aerial platform allowed a regional scale canopy arthropod diversity study with many replicates using a direct sampling technique; 2) the results demonstrated differences between canopy and understorey spider diversity and the complementary nature of the fauna in these two habitats; 3) results also showed that spider assemblage composition differs between sugar maple and American beech canopies, and that different processes drive spatial patterns of diversity in the canopy and the understorey. This original paper should inspire further canopy athropod studies in temperate forests because of the possibilities offered by this new canopy access method and the demonstration that the canopy spider fauna is relevant to the biological diversity of north-temperate hardwood forests. Chapter 4 This original paper is the first investigation of the ballooning propensity of spider species across many families in a forest and in the canopy. The adaptive links among ballooning propensity, life history traits and habitat conditions make ballooning an important research area. The knowledge gained in this study can be applied to other areas of ecology like understanding the processes that account for spatial patterns
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