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Patterns and Drivers of Terrestrial Arthropod Biodiversity in Northern Canada

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Patterns and Drivers of Terrestrial Arthropod Biodiversity in Northern Canada Patterns and drivers of terrestrial arthropod biodiversity in northern Canada Crystal M. Ernst Department of Natural Resource Sciences McGill University Montreal, Quebec, Canada April 2015 A thesis submitted to McGill University in partial fulfillment of the requirements of the degree of Doctor of Philosophy © Crystal M. Ernst 2015 Abstract The overarching goal of this thesis was to describe patterns of terrestrial arthropod biodiversity and community structure in northern Canada, and to explore the underlying drivers and mechanisms that are responsible for these patterns. The term “biodiversity” is used here in a broad sense that includes both taxonomic (TD) and functional (FD) diversity. Ground-dwelling arthropods, especially beetles (Coleoptera), were used as model taxa, and were collected using standardized methods from twelve locations in the three northernmost ecoclimatic zones of Canada. Beetle biodiversity changes over time and space. Over the course of one active season, rapid species and functional turnover were observed in two major habitats in one subarctic location (Kugluktuk, Nunavut). While some functional groups were apparent only for brief periods of time, entomophagous predators consistently dominated the assemblage structure in biomass and abundance. This dominance by carnivores was observed consistently throughout the study, regardless of spatial or taxonomic scope. This inverted trophic structure suggests that predators may rely on alternative, non-epigeic prey items. A natural history study of previously unknown host-parasite interactions between beetles and nematomorphs (Gordionus n. sp.) suggests that beetles use alate insects with aquatic larval stages as an important nutrient subsidy. Across the entire study region, beetle TD and FD, as well as overall assemblage structure, display strong negative relationships with latitude, which conforms to the classical latitudinal gradient of diversity. After considering many spatial, environmental and climatic variables, the most significant driver of beetle biodiversity and assemblage structure over time and space was climate, particularly temperature. i When the taxonomic scope of the research was expanded to include all ground-dwelling macroarthropod taxa, and the functional scope refined with a multidimensional functional trait- based approach, the same patterns and processes were observed. Additionally, it was found that functional redundancy was greater in the high arctic than in the warmer ecoclimatic zones further south, despite a paucity of taxa in the high arctic. This supports the hypothesis that environmental constraints are more important in regions with harsh climates, and play a greater role in diversity and community assembly processes than niche differentiation. The biodiversity (TD and FD) and structure of terrestrial macroarthropod communities in northern Canada have several consistent patterns: (1) large-scale negative relationships between biodiversity/assemblage structure and latitude; (2) strong correlations between TD and FD; and (3) the dominance of active predatory taxa. The most important finding of this study is that climate (temperature) gradients provide the best explanation for the variability observed in arthropod biodiversity and assemblage structure over time and across space. Lastly, the effects of climate on biodiversity and community assembly seem to be more pronounced in the high arctic than in more southerly biomes. Given the rapid and significant rise in temperature projected for northern biomes and the fact that predatory taxa are often more sensitive to changes in their environments, major changes to arthropod diversity are expected in the north, with implications for the stability of northern ecosystems. ii Résumé L'objectif principal de cette thèse est de décrire les patrons de biodiversité des arthropodes terrestres, la structure de leurs communautés dans le Nord du Canada, et d'explorer les forces et les mécanismes responsables de ces patrons. Le terme «biodiversité» est utilisé ici dans un sens large qui comprend à la fois la diversité taxonomique (DT) et fonctionnelle (DF).Les arthropodes vagabonds, en particulier les coléoptères (Coleoptera), utilisés comme taxon modèle, ont été recueillies à l'aide de méthodes normalisées dans douze sites répartis dans les trois zones écoclimatiques les plus septentrionales du Canada. La biodiversité de coléoptères varie dans l’espace et le temps. Au cours d'une saison active, un changement taxonomique et fonctionnel rapide a été observé dans les deux habitats étudiés à Kugluktuk, Nunavut, dans la région subarctique. Alors que certains groupes fonctionnels n’étaient observés que pour de brèves périodes de temps, les prédateurs entomophages dominaient constamment la structure de l’assemblage en biomasse et en abondance. La prépondérance des carnivores en Arctique a été observée régulièrement pendant toute l'étude, quelle que soit la portée spatiale ou taxonomique. Cette structure trophique inversée suggère que les prédateurs pourraient se nourrir de proies non vagabondes. Une étude de l'histoire naturelle des interactions hôte-parasite précédemment inconnues entre les coléoptères et les nématomorphes (Gordionus n. Sp.) suggère que les coléoptères consomment des insectes ailés avec des stades larvaires aquatiques comme une source importante de nutriments. À travers toute la région d'étude, la DT et la DF des coléoptères, ainsi que la structure globale d'assemblage, étaient négativement corrélées avec la latitude, ce qui est conforme au gradient latitudinal classique. Après avoir examiné de nombreuses variables spatiales, environnementales et climatiques, la force qui semble influencer le plus la biodiversité et la iii structure de l’assemblage des coléoptères au Nord, et ce, dans l’espace ou dans le temps, est le climat, en particulier la température. Lorsque tous les taxons de macroarthropodes terrestres vagabonds ont été inclus, et que la variabilité fonctionnelle est décrite à l’aide de plusieurs traits fonctionnels, les patrons de biodiversité et de structure observés ainsi que les forces sous-jacentes étaient similaires. La redondance fonctionnelle était plus grande dans l'Extrême-Arctique que dans les zones écoclimatiques plus au sud, et ce, en dépit d'un manque de taxons dans l'Extrême-Arctique. Cela confirme l'hypothèse que les contraintes environnementales sont plus importantes dans les régions au climat rigoureux, et jouent un rôle plus important dans la variation de diversité et d'assemblage que la force de différenciation de niche écologique. La biodiversité (DT et DF) et la structure des communautés de macroarthropodes terrestres dans le Nord du Canada ont plusieurs patrons compatibles: (1) relations négatives à grande échelle entre la biodiversité / la structure d'assemblage et la latitude; (2) de fortes corrélations entre DT et DF; et (3) la domination des taxons prédateurs actifs. La conclusion la plus importante de cette étude est que le climat (température) semble être la force qui explique le mieux la variabilité observée dans la biodiversité des arthropodes et la structure d’assemblage dans le temps et dans l'espace. Enfin, les effets du climat sur la biodiversité et l'ensemble de la communauté semblent être plus prononcés dans l'Extrême-Arctique que dans les biomes plus au sud. Compte tenu de la hausse rapide et importante de la température prévue pour les régions nordiques et le fait que les taxons prédateurs sont souvent plus sensibles aux changements dans leur environnement, des changements majeurs de la diversité des arthropodes sont attendus dans le nord, avec des implications importantes pour la stabilité des écosystèmes nordiques. Translation: Sarah Loboda iv Table of Contents Abstract ........................................................................................................................................... i Résumé .......................................................................................................................................... iii Table of Contents .......................................................................................................................... v List of Tables ................................................................................................................................ ix List of Figures .............................................................................................................................. xii Thesis format, contributions of authors, and contributions to knowledge ........................... xvi Acknowledgments .................................................................................................................... xviii Chapter 1: Introduction, literature review, and objectives ...................................................... 1 1.1 Introduction ........................................................................................................................... 1 1.2 Literature Review .................................................................................................................. 3 1.2.1 Patterns of terrestrial diversity ........................................................................................ 3 1.2.2 A paradigm shift in ecology ........................................................................................... 5 1.2.3 Mechanisms underlying biodiversity patterns ................................................................ 9 1.2.4 Biodiversity in northern ecosystems: change
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