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Impacts of Sedimentation on the Structure And IMPACTS OF SEDIMENTATION ON THE STRUCTURE AND FUNCTIONING OF AGRICULTURAL STREAM COMMUNITIES A thesis submitted in partial fulfilment of the requirements for the Degree of Doctor of Philosophy in Ecology in the University of Canterbury by Francis J. Burdon University of Canterbury 2013 Deputy Vice-Chancellor’s Office Postgraduate Office Co-Authorship Form This form is to accompany the submission of any thesis that contains research reported in co- authored work that has been published, accepted for publication, or submitted for publication. A copy of this form should be included for each co-authored work that is included in the thesis. Completed forms should be included at the front (after the thesis abstract) of each copy of the thesis submitted for examination and library deposit. Please indicate the chapter/section/pages of this thesis that are extracted from co-authored work and provide details of the publication or submission from the extract comes: Chapter 1: “Sedimentation and stream ecosystems: a review of impacts on community structure and functioning” by Francis J. Burdon and Jon S. Harding has been submitted for review and comments have been received. Chapter 2: “Habitat loss drives threshold response of benthic invertebrate communities to deposited sediment in agricultural streams” by Francis J. Burdon, Angus R. McIntosh, and Jon S. Harding is in press in the journal “Ecological Applications” Please detail the nature and extent (%) of contribution by the candidate: In Chapter 1, the data analysis and writing were largely performed by the candidate as lead author (95% of total contribution). In Chapter 2, the data analysis and writing were largely performed by the candidate as lead author (90% of total contribution) Certification by Co-authors: If there is more than one co-author then a single co-author can sign on behalf of all The undersigned certifys that: . The above statement correctly reflects the nature and extent of the PhD candidate’s contribution to this co-authored work . In cases where the candidate was the lead author of the co-authored work he or she wrote the text Name: Jon S. Harding Signature: Date: 07/05/2013 Ad excelsiora TABLE OF CONTENTS Acknowledgements 2 Thesis Abstract 4 Preface 6 Chapter 1 12 Sedimentation and stream ecosystems: a review of impacts on community structure 12 and functioning Chapter 2 53 Habitat loss drives threshold response of benthic invertebrate communities to 54 deposited sediment in agricultural streams Chapter 3 91 Niche-space compression influenced by sedimentation in agricultural stream food 92 webs Chapter 4 138 Sedimentation leads to cascading effects on biodiversity, consumer-resource 140 interactions, and ecosystem function in agricultural streams Chapter 5 180 Synthesis: sedimentation and the structure and functioning of stream communities 180 Appendices 224 Chapter 2: Appendices A-H 224 Chapter 3: Appendices A-D 250 Chapter 4: Appendices A-D 272 1 ACKNOWLEDGEMENTS ACKNOWLEDGEMENTS I would like to thank my senior supervisor, Jon Harding and associate supervisor, Angus McIntosh for the opportunity to conduct doctoral research at the University of Canterbury. The funding provided by the Mackenzie Charitable Foundation and the Brian Mason Scientific and Technical Trust has been indispensable, and I laud those organisations for the services they are providing to the Canterbury region. I would like to thank the Canterbury Branch of the Royal Society of New Zealand, the SIL Trust, and the Society for Freshwater Sciences Endowment Committee for the generous travel funding each of these groups provided to me during my doctoral research. Enabling early career researchers from New Zealand to travel to overseas conferences is an immensely valuable service, and I am very grateful for the financial support. I greatly appreciate the excellent assistance provided by Hayley Stoddard in the laboratory. Your skills are indispensable, and I sincerely hope your valuable contributions to the research process are justly rewarded. I appreciate the help Tom Moore provided sorting leaf packs. I thank those that assisted with fieldwork, including Richard White, Nixie Boddy, Duncan Grey, Danladi Umar, and Steve Pohe. Technical assistance provided by Linda Morris, Jan MacKenzie, Milan Marinov, Nick Etheridge, and Alan Woods was much appreciated. I would like to thank those that provided useful discussions regarding a variety of statistical and scientific issues, including Tanya Blakely, Phil Jellyman, Pete McHugh, Jon O’Brien, Elizabeth Graham, Kristy Hogsden, Dev Niyogi, and Kevin Simon. I greatly appreciate the patient souls who provided helpful comments on my written work, including the erudite Mike Winterbourn and my father, Rowland Burdon. 2 ACKNOWLEDGEMENTS I would like to thank those past and present who have supported the School of Biological Sciences Journal Club. It is a beacon of intellectual verve and collegiality that I sincerely hope continues to burn bright into the future. It would be remise of me not to highlight the contribution of Ant Poole and Will Godsoe to this valuable institution. I would also like to thank those academics whose foresight and contacts enabled eminent scientists from overseas, such as Joel Cohen, Dominique Gravel, and Spencer Barrett, to give memorable talks at the university that made a lasting impression. I greatly appreciate those involved with New Zealand Natural Sciences Journal for their contributions over the years. My service to this worthy publication has been at times a challenging experience, but most rewarding and I sincerely hope to see the journal continue on well into the future. I have to thank Eunice McKessar from IES Christchurch for one of the best part-time jobs a PhD student could hope for, and acknowledge the inspirational students from all over the United States who attended the University of Canterbury program during my employment. I thank my fellow students at the University, including those in the FERG group for the good times, whether it came hiking trails, playing soccer, cooking (and eating) feasts, or just sitting outside on the grass and talking. I appreciate those friends who I have seen recently for their good company, and apologise to the ones who I have neglected during the rigours of the PhD process. To my long suffering flatmates, I sincerely hope to compensate for my absence in the last few months. Finally, I would especially like to thank my parents, whose love and support has been unwavering over the years. Your confidence has helped to propel me towards achieving my goals. 3 THESIS ABSTRACT THESIS ABSTRACT The excessive deposition of fine inorganic sediment (<2 mm) is a major pathway by which agricultural land uses exert pressure on stream ecosystems. However, less well understood are the underlying mechanisms driving threshold biotic responses and the ecological consequences of community changes to sedimentation. Reviewing the literature, I found that sedimentation can affect algal and detrital pathways, and invertebrate community composition may show abrupt shifts with increased sediment. Moreover, functional changes to communities potentially leads to simpler food webs, with altered interactions and decreased ecosystem function. After identifying these knowledge gaps, I conducted survey and experimental research using agricultural streams on the Canterbury Plains of New Zealand’s South Island. Results from my survey of 30 streams along a sedimentation gradient showed that pollution-sensitive invertebrates (% EPT; Ephemeroptera, Plecoptera, Trichoptera) demonstrated threshold responses to sediment that varied with spatial scale, and change-point analysis indicated marked declines beyond 20% fine sediment covering streambed reaches. Structural equation modeling indicated that decreased habitat availability was a key mechanism contributing to these changes. To better understand the functional consequences of altered community structure, I investigated food webs in 12 streams along the gradient. The results showed a compression of community trophic niche space, suggesting that in particular, primary consumers became trophically more equivalent. The simplification of stream food webs with increasing sediment appeared to be the result of functional changes to invertebrate communities, with fewer specialised consumers, and shifts in the availability of basal resources. Using field and laboratory experiments investigating litter breakdown and invertebrate feeding, I found that the net consequence of functionally less diverse stream communities with increased sediment was impaired ecosystem function, demonstrated by 4 THESIS ABSTRACT a reduction in litter breakdown rates. The reduction of detrital resource availability through burial by sediment in laboratory mesocosm experiments strongly influenced detrital consumption rates, thus leading to reduced growth and survival of detritivorous caddisflies. The survey and experimental results support my postulate that sediment deposition causes environmental stress by degrading benthic habitat and making associated food resources (e.g., periphyton and leaves) less available. Overall, my results have provided new insights into sediment impacts on stream communities and have furthered our understanding of how these changes affect the structure and functioning of stream ecosystems. 5 PREFACE PREFACE This thesis addresses the potential influences of stream sedimentation at multiple levels of biological organisation (e.g., communities, food webs, and ecosystems). I postulated
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