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THE ROLE OF CHIRONOMIDS AS PALEOECOLOGICAL INDICATORS OF EUTROPHICATION IN SHALLOW LAKES ACROSS A BROAD LATITUDINAL GRADIENT by Emily Meaghan Stewart A thesis submitted to the Department of Biology In conformity with the requirements for the degree of Doctor of Philosophy Queen’s University Kingston, Ontario, Canada (January, 2018) Copyright © Emily Meaghan Stewart, 2018 Abstract The aquatic larvae of chironomids (Diptera, Chironomidae) were historically classified according to lake trophic status, and taxa classified as “eutrophic” were labeled as such because of adaptations for surviving hypoxic or anoxic conditions in the hypolimnion of stratified eutrophic lakes. As such, sedimentary chironomid assemblages have been used to reconstruct production-related variables (nutrients, chlorophyll-a), though this has been problematic, especially in shallow systems, because the response of chironomids to eutrophication is mediated through secondary environmental gradients including oxygen concentration, habitat, and food quality/quantity. In this thesis, eutrophic sewage ponds in the Canadian High Arctic were used to demonstrate that oxygen, not nutrients, is the primary control of chironomid species assemblages. The ability to explicitly test the influence of oxygen versus nutrients on chironomid distributions was made possible by the 24-hr daylight (continuous photosynthesis) and shallow, wind-mixed water column that resulted in oxygen concentrations that were decoupled from the effects of elevated nutrients and production. The subfossil chironomid assemblages were complacent during historical eutrophication, in contrast to marked changes in diatom assemblages, which have a direct physiological relationship with nutrients. Similarly, in shallow eutrophic ponds on islands in Lake Ontario, chironomid assemblages did not appear to be governed by the large gradient in total phosphorus due to the presence or absence of waterbird nesting colonies, but rather by habitat and possible bird- mediated heavy metal pollution. In a subarctic lake that was formerly used for sewage disposal, chironomid assemblages were relatively unresponsive to eutrophication in comparison with the larger turnover in diatom species. However, periods of low oxygen observed in the temperate and subarctic sites may explain the higher (but still low) relative abundances of hypoxia-tolerant Chironomus species at these sites compared to the High Arctic ponds. Together, this research demonstrates the problems associated with classifying chironomids based on nutrient levels. Since it is uncommon to examine chironomid responses to eutrophication across latitude, this thesis offers a relatively unstudied perspective ii of chironomid ecology, emphasizing that some of the assumptions of temperate chironomid ecology (with regards to eutrophication) may not necessarily hold true when applied at higher latitudes. iii Co-Authorship Chapter 2 was co-authored by Reba McIver, Neal Michelutti, Marianne S.V. Douglas, and John P. Smol. I collected 2011 samples, co-mentored Reba McIver with Neal Michelutti (supervised by John Smol), analysed chironomids, completed data analysis (including statistical work), and was the primary author of the paper. Reba McIver identified diatoms under the tutelage of Neal Michelutti. Marianne Douglas led 2011 field work, and together with John Smol, oversaw study design. Previous field seasons were also led by Marianne Douglas and John Smol. This chapter was published separately: Stewart EM, McIver R., Michelutti N, Douglas MSV, Smol JP (2014) Assessing the efficacy of chironomid and diatom assemblages in tracking eutrophication in High Arctic sewage ponds. Hydrobiologia 721:251–268 Chapter 3 was co-authored by Neal Michelutti, Christopher Grooms, Linda E. Kimpe, Jules M. Blais, and John P. Smol. I participated in study design and field work; analysed chironomids, sedimentary chlorophyll-a, and water chemistry; performed statistical analyses; and was the primary author of the chapter. Neal Michelutti aided with data analysis and interpretation, as well as (with Christopher Grooms) completed field work. Linda Kimpe and Jules Blais completed elemental analysis on the cores, and aided with the interpretation of core radiochronologies. John Smol oversaw study design and data interpretation. Chapter 4 was co-authored by Kathryn E. Hargan, Branaavan Sivarajah, Linda E. Kimpe, Jules M. Blais, and John P. Smol. I participated in study design and field work, processed for chironomids and chlorophyll-a, did statistical analyses, and was the primary author of the paper. Kathryn Hargan completed sterol analysis and interpretation under the supervision of Jules Blais. Branaavan Sivarajah completed diatom analysis. Jules Blais led field work and aided in sterol analysis. Linda Kimpe was responsible for core radiochronology. John Smol oversaw project design. The manuscript for this chapter is under review in the journal Arctic (submission number: 17-134). Appendix A was co-authored by Kathryn E. Hargan, Neal Michelutti, Christopher Grooms, Linda E. Kimpe, Mark L. Mallory, John P. Smol, and Jules M. Blais. I helped with study design, collected and analysed 210Pb data, analysed total lead and stable lead isotope data, and was the second author of the iv paper. Kathryn Hargan completed sterol analysis and was first author of the manuscript. Neal Michelutti and Christopher Grooms did field work. Linda Kimpe and Jules Blais oversaw sterol data analysis and interpretation, as well as collected total lead and stable lead isotope data. Mark Mallory helped with data analysis with respect to birds. John Smol aided in project concept design. Data were originally included as supplementary material in: Hargan KE, Stewart EM, Michelutti N, Grooms C, Kimpe L, Mallory M, Smol JP, Blais JM (in review) Incorporating sterols and stanols as biomarkers for tracking waterbird impacts to temperate ponds. Proc R Soc Lond B Biol Sci: manuscript ID RSPB-2017-2669 Appendix B was co-authored by Neal Michelutti, Mina Vu, Christopher Grooms, Linda E. Kimpe, John P. Smol, and Jules M. Blais. I took part in study design, co-mentored Mina Vu with Neal Michelutti (supervised by John Smol), analysed data, performed statistical analyses, and was the primary author of the manuscript (in prep.). Mina Vu analysed diatoms with Neal Michelutti. Christopher Grooms did field work and collected chlorophyll-a data. Linda Kimpe and Jules Blais performed stable isotope analysis and helped with interpretation. John Smol partook in study design and overseeing data analysis and interpretation. v Acknowledgements My sincerest thanks to my supervisor, John Smol, and co-supervisor, Marianne Douglas, without whom I would not be a paleolimnologist. John, thank you for taking me in as a 3rd year summer student and letting me stay for a defining 7 years of my life. You both fostered my love of limnology and introduced me to one of my favourite things: The Arctic. Thank you for this opportunity, and thank you for sharing your infectious passion for learning and exploring with your students. I could not have had a better start to my path as a scientist. Thank you to my committee members, Drs. Shelley Arnott and Scott Lamoureux, for positive committee meeting experiences that churned out better ideas and led to stronger work. Thank you to PEARL, current and former, you have always been a family to me. PEARL was and is full of my best friends - a support network and a collection of wonderful minds that I have the privilege to know. I would like to express an immense amount of gratitude to Neal Michelutti – you were stuck with me from the beginning and dealt with the terrible writing and numerous questions! Thank you for being a superior role model in science writing and interpretation; your patience and constructive feedback have always been helpful. I cannot name everyone who has made my time here memorable, but needless to say, if you were or are a part of PEARL, you are on that list! Finally, thank you to the Secchi table for consistently hosting some of the very best and funniest experiences I’ve ever had, as well as the most productive science conversations – especially during coffee party. Thank you to my family, especially my poor parents and sister, who withstood a large portion of my graduate woes without (much) complaint. You are more important than I could ever say, and you deserve at least one of the letters of this PhD (the “P” perhaps). An enormous thank-you is awarded to my husband, Graham, you obviously deserve the other letters. Not only did you come into my life at the beginning of my Master’s, but a) you decided to stay with me through the PhD, despite the stress and craziness, b) you are absolutely my favourite person, and c) you gave me an amazing and supportive second family, as well. The love and support of my family through this was and still is immeasurable. vi Table of Contents Abstract ......................................................................................................................................................... ii Co-Authorship.............................................................................................................................................. iv Acknowledgements ...................................................................................................................................... vi List of Figures .............................................................................................................................................