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Thesis Submitted to the Department of Biology The effects of habitat connectivity and regional heterogeneity on artificial pond metacommunities by Michael Torrance Pedruski A thesis submitted to the Department of Biology in conformity with the requirements for the degree of Master of Science Queen’s University Kingston, Ontario, Canada October, 2008 Copyright © Michael Torrance Pedruski, 2008 Abstract While much evidence suggests that ecosystem functioning is closely related to biodiversity, present rates of biodiversity loss are high. With the emergence of the meta- community concept ecologists have become increasingly aware that both local processes (e.g. competition, predation), and regional processes (e.g. dispersal and regional hetero- geneity) affect ecological communities at multiple spatial scales. I experimentally inves- tigated the effects of habitat connectivity and regional heterogeneity on biodiversity, community composition, and ecosystem functioning of artificial pond metacommunities of freshwater invertebrates at the local (α), among-community (β), and regional (γ) spa- tial levels. There was a significant effect of habitat connectivity on mean local richness, but mean local Simpson diversity, mean local functional diversity (FD), and all the three in- dices of ecosystem functioning investigated (regional abundance, invertebrate biomass, and chlorophyll a concentration) were unaffected by connectivity levels. Regional het- erogeneity had no effect on local diversity, but enhanced both among-community rich- ness and among-community Simpson diversity. Conversely, connectivity reduced among- community Simpson diversity. All indices of regional diversity were unaffected by either connectivity or heterogeneity. Despite expectations that there would be strong interac- tions between the effects of connectivity and heterogeneity on species richness, there were no interactions for any index of biodiversity at any spatial scale. ii Invertebrate community composition was unaffected by either connectivity or heterogeneity, though there was a significant effect of heterogeneity on its variance. Nei- ther connectivity nor heterogeneity had significant effects on any index of ecosystem functioning, nor among-community coefficients of variation of ecosystem functioning. Connectivity appears to act mainly as a force homogenizing habitat patches in a region, as opposed to having strong effects in and of itself on communities. Conversely, heterogeneity acts largely as a diversifying force, maintaining differences between com- munities within a region, but, similar to connectivity, it does not have clear effects on communities at the local scale. Despite the different processes expected to act in homo- geneous and heterogeneous regions, it does not appear that connectivity and heterogene- ity interact strongly. iii Co-Authorship This thesis conforms to the manuscript format as outlined by the School of Graduate Studies and Research. The manuscript that is a direct result of this thesis and its coauthors is: Pedruski, M. T., and S.E. Arnott. In preparation for Ecology. The effects of habitat con- nectivity and regional heterogeneity on biodiversity, community composition, and ecosystem functioning of artificial pond metacommunities. iv Acknowledgments First and foremost I owe an inestimable debt of gratitude to my supervisor and friend Shelley Arnott whose guiding hand not only contributed greatly to the develop- ment of this thesis, but also to my personal development as a scientist and as an individ- ual. Many others contributed thoughtful advice and criticism that greatly improved the quality of this research. I’m grateful to Angela Strecker, Alison Derry, Carol Frost, Shan- non MacPhee, and Leah James for suggestions regarding experimental design and analy- sis. Further gratitude is due to the members of my supervisory committee, Lonnie Aarssen and Bruce Tufts who contributed both appreciated insights and resources to the execution of my project. A study of this magnitude could not have been executed by one person, and I would be remiss to not thank all of those who helped out in the field or in the lab: Hannah Kent, Colin McCleod, Megan MacLennan, Theresa Patenaude, Angela Strecker, Justin Shead, my parents, Catherine Gieysztor, Alexandra Howard, Steve Hawrylyshyn, Janine Lee, and Niki Willie. Special thanks are due to the last three, Steve for sticking it out the whole four months with me through good times and bad, Janine for tolerating everything I threw at her, supporting me 110%, and always putting a smile on my face; and Niki for doing an inordinate amount of work out of the sheer goodness of her heart, as well as providing inspiration to climb trees. To the staff of QUBS, Frank Phelan and Floyd Conor, without whose logistical support this study would have been impossible, I am grateful. v To all the members of the Arnott lab with whom I have had the privilege to work, you made it worth while coming into the office on a daily basis: Johanna Pokorny, Lisa Duke, Esther Chan, Megan MacLennan, Theresa Patenaude, Darryl McGrath, Corinne Daly, Liz Hatton, Justin Shead, Colleen Inglis, Derek Gray, Anneli Jokela, Angela Strecker, Alison Derry, Shannon MacPhee, and Leah James. Finally, I was supported by an NSERC CGS-M award and PGS-M extension for the duration of this research, for which I am grateful. vi Table of Contents Abstract ...............................................................................................................................ii Co-authorship .....................................................................................................................iv Acknowledgments ...............................................................................................................v Table of Contents ..............................................................................................................vii List of Tables ......................................................................................................................ix List of Figures .....................................................................................................................x Chapter 1 - General Introduction ....................................................................................1 The Metacommunity Concept ....................................................................................4 Biodiversity in Metacommunities ..............................................................................6 Effects of Connectivity and Heterogeneity in Single Species Systems ......................8 Competitive Coexistence in Metacommunities ..........................................................9 Trophic Interactions in Metacommunities ...............................................................11 Ecosystem Functioning and Metacommunities .......................................................13 Conclusion on Effects of Connectivity and Heterogeneity on Metacommunities ....15 Freshwater Invertebrates as a Model System ..........................................................15 Thesis Objectives .....................................................................................................18 Chapter 2 - The effects of habitat connectivity and regional heterogeneity on biodiversity, community composition, and ecosystem functioning of artificial pond metacommunities ............................................................................................................19 Abstract ....................................................................................................................20 Introduction .............................................................................................................21 Methods ...................................................................................................................25 Experimental Design .................................................................................25 Experimental Set-Up .................................................................................25 Experimental Treatments ..........................................................................26 Sampling Protocol .....................................................................................27 Invertebrate Enumeration and Identification ...........................................28 Community Diversity ................................................................................30 Habitat Separated Analyses ......................................................................32 Community Composition ..........................................................................32 Biomass Calculations ...............................................................................32 Data Analysis ............................................................................................33 Results .....................................................................................................................35 Environmental Variables ...........................................................................35 Community Diversity ................................................................................35 Habitat Separated Analyses ......................................................................37 Community Composition ..........................................................................37 Abundance, Biomass, Chlorophyll a .........................................................38 Relationship between Diversity and Ecosystem Functioning
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