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Investigations Into Food Web Structure in the Beaufort Sea Investigations into food web structure in the Beaufort Sea by Ashley Stasko A thesis presented to the University of Waterloo in fulfillment of the thesis requirement for the degree of Doctor of Philosophy in Biology Waterloo, Ontario, Canada 2017 © Ashley Stasko 2017 Examining committee membership The following served on the Examining Committee for this thesis. The decision of the Examining Committee is by majority vote. External Examiner Dr. Eddy Carmack Research Scientist (Fisheries and Oceans Canada) Supervisor(s) Dr. Michael Power Professor Dr. Heidi Swanson Assistant Professor, University Research Chair Internal Member Dr. Roland Hall Professor Internal-external Member Dr. Simon Courtenay Professor, Canadian Water Network Scientific Director Other Member(s) Dr. James Reist Research Scientist (Fisheries and Oceans Canada) ii Author’s declaration This thesis consists of material all of which I authored or co-authored: see Statement of Contributions included in the thesis. This is a true copy of the thesis, including any required final revisions, as accepted by my examiners. I understand that my thesis may be made electronically available to the public. iii Statement of contributions While the research in this thesis was my own, co-authors provided valuable input and contributions to each chapter. The research was conducted in association with the Beaufort Regional Environmental Assessment Marine Fisheries Project (BREA MFP). Fisheries and Oceans Canada oversaw the design and execution of the field sampling program, led by Andrew Majewski and James Reist. A full list of field participants, including myself, can be found in Majewki et al. (2016). Over-arching research objectives and themes for the PhD were developed collaboratively by myself, Michael Power, Heidi Swanson, and James Reist in concert with priorities defined by the BREA MFP. Writing, data analysis, and specific study design ideas for each chapter were primarily developed by myself, with contributions from Michael Power and Heidi Swanson (University of Waterloo), as well as Bodil Bluhm (UiT – the Arctic University of Norway). Specific additional contributions to each chapter from co-authors are outlined below. Chapter 2: Influences of depth and pelagic subsidies on the size-based trophic structure of Beaufort Sea fish communities. Ashley D. Stasko, Heidi Swanson, Andrew Majewski, Sheila Atchison, James Reist, and Michael Power. Aside from the co-author contributions stated above, AM provided fish distribution data and SA provided biomass and trawl area calculations. All authors provided significant edits and helpful insights during manuscript preparation. The text presented in this thesis is a pre-copy editing version of a manuscript published as: Stasko AD, Swanson H, Majewski A, Atchison S, Reist J, Power M (2016) Influences of depth and pelagic subsidies on the size-based trophic structure of Beaufort Sea fish communities. Marine Ecology Progress Series, 549: 153-166, doi: 10.3354/meps11709 iv Chapter 3: Benthic-pelagic trophic coupling in an Arctic marine food web along gradients of water mass structure and organic matter input. Ashley D. Stasko, Bodil A. Bluhm, Heidi Swanson, Christine Michel, Philippe Archambault, Andrew Majewski, James D. Reist, and Michael Power. Aside from the co-author contributions stated above, BAB provided significant input regarding study design and results, and acted as a primary advisor during a period of research abroad. CM provided algal biomass and primary productivity data. PA provided analyses of sediment granulometry, organic matter content, and pigment concentrations. All authors provided significant edits and helpful insights during manuscript preparation. This chapter has been submitted for publication under the same title and authorship in Marine Ecology Progress Series (manuscript ID 22559). Chapter 4: Relationships between depth and δ15N of Arctic benthos vary among regions and trophic functional groups. Ashley D. Stasko, Bodil A. Bluhm, James D. Reist, Heidi Swanson, and Michael Power. Aside from the co-author contributions stated above, all authors provided significant edits and helpful insights during manuscript preparation. This manuscript has been submitted for publication under the same title and authorship in Deep Sea Research I: Oceanographic Research Papers (manuscript ID DSRI_2017_43). Chapter 5: Responses of benthic functional food web structure to variable food supply in two Arctic shelf ecosystems. Ashley D. Stasko, Shannon MacPhee, Philippe v Archambeault, Jane Eert, Andrew Majewski, James Reist, Heidi Swanson, and Michael Power. Aside from the co-author contributions stated above, SM provided samples for epifauna and infauna. PA provided analyses of sediment granulometry, organic matter content, and pigment concentrations. JE provided oceanography data. vi Abstract The effects of climate change on marine ecosystems are most pronounced in the Arctic, where ice free summers have been predicted to occur by mid-century. Climate- related changes to sea ice phenology, oceanographic habitat characteristics, and primary production regimes will likely have strong effects on ecosystem structure that could alter energy pathways, species distributions, food web dynamics, and secondary production. Knowledge of many offshore Arctic ecosystems remains poor, undermining the ability to predict the effects of a changing climate on food web structure and function. This thesis capitalizes on the first comprehensive offshore sampling program in the Canadian Beaufort Sea and Amundsen Gulf to address substantial knowledge gaps regarding fish and invertebrate food web structure in the region. Trophic structure and benthic-pelagic linkages for biological communities on the continental shelf and slope were examined using stable isotope values measured in 127 fish and invertebrate taxa, biomass distributions, and a database of biological functional traits compiled for 166 taxa. Four empirical studies were conducted to test hypotheses regarding the responses of trophic structure to environmental gradients of depth, organic matter input regimes, water mass structure, and benthic food supply. Understanding food web structure and its link to large-scale environmental gradients will be key to assessing and predicting the effects of climate change on offshore marine communities in the Canadian Beaufort Sea and Amundsen Gulf. In Chapter 2, benthic-pelagic coupling via active biological transport was identified as important for sustaining fish communities in the Beaufort Sea. Lower availability of benthic resources with increasing depth restricted biomass production for small size classes of fish in deep habitats. In those same fish communities, pelagic subsidies obtained by benthopelagic fishes were important for maintaining a relatively vii high biomass of large-bodied fish in deep habitats. When fish and invertebrates were considered together in Chapter 3, benthic-pelagic coupling weakened eastward alongshore, across three regions. Benthic-pelagic coupling was (1) highest west of the Mackenzie River where sinking flux of pelagic particulate organic matter (POM) is known to be relatively high, (2) intermediate on the Mackenzie Shelf where riverine inputs of terrestrial organic matter dominate the sediment, and (3) lowest in the Amundsen Gulf where strong pelagic grazing is known to limit POM sinking flux to the benthos. Within all regions considered, benthic-pelagic coupling was consistently weakest in slope habitats underlying the transition between Pacific- and Atlantic-origin waters, where much of the organic carbon is transformed or intercepted in the water column. Analyses in Chapter 4 indicated that the dominance of terrestrial POM discharged from the Mackenzie River in the Beaufort Sea dampened depth-related changes in the δ15N values of suspension/filter feeders, infaunal deposit feeders, and bulk sediment. In contrast, a faster rate of change in consumer and sediment δ15N with depth was observed in the Amundsen Gulf. Relatively high primary production in the Amundsen Gulf likely promoted intensified biological transformation of autochthonous POM in the pelagic zone and lower downward POM flux, causing greater change in POM δ15N. Surprisingly, when isotopic diversity was weighted by species biomasses in Chapter 5, most benthic communities in the Canadian Beaufort Sea and Amundsen Gulf were found to rely on similarly diverse ranges of sedimentary organic matter, regardless of the sources. Trait-based functional diversity indicated that shelf edge communities maintained a relatively high diversity of biological trophic traits, presumably to exploit pulsed food inputs associated with dynamic shelf break hydrography. Several lines of evidence supported a role for episodic food inputs in structuring shelf edge trait composition. However, pairwise relationships between viii trophic traits and indicators of benthic food supply were not significant at the regional scale. Functional redundancy was low across most of the region, suggesting benthic food web function will be sensitive to species loss. The research in this thesis presents the first comprehensive empirical studies of benthic food web structure for offshore fish and invertebrate communities in the Canadian Beaufort Sea and Amundsen Gulf. Each study proposes causal explanations for spatial patterns in food web structure based on data for habitat characteristics, species biomass distributions, and previously documented physical and biological properties of the regions.
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