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On Stream Ecosystem Functioning and Structure in Cape Breton Highlands, Nova Scotia

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On Stream Ecosystem Functioning and Structure in Cape Breton Highlands, Nova Scotia EFFECTS OF HYPERABUNDANT MOOSE (Alces alces) ON STREAM ECOSYSTEM FUNCTIONING AND STRUCTURE IN CAPE BRETON HIGHLANDS, NOVA SCOTIA by © Jessica MacSween A thesis submitted to the School of Graduate Studies in partial fulfilment of the requirement for the degree of Master of Science (Environmental Science) Department of Biology Memorial University of Newfoundland January 2018 St. John’s Newfoundland and Labrador ABSTRACT Terrestrial consumers can influence forest regeneration, but few studies have investigated the impacts of terrestrial consumers on freshwater ecosystems. I investigated the potential for cross-ecosystem effects of hyperabundant moose on stream ecosystems in the Cape Breton Highlands, Nova Scotia. I predicted that watersheds with higher moose impacts would have higher stream temperatures, total nitrogen, electrical conductivity, periphyton biomass, and macroinvertebrate abundances. I analyzed existing long-term data on 3rd and 4th order streams and conducted a field study on 1st and 2nd order streams to test for cross- ecosystem impacts of moose in the Cape Breton Highlands. I found evidence for moose impacts on total nitrogen and electrical conductivity. However, I found no evidence for higher stream temperatures and periphyton biomass, with limited evidence of moose impacts on macroinvertebrate abundances. This study provides insight into the effects of large ungulates within and across boreal forest ecosystems with potential implications for landscape-scale management of hyperabundant ungulates. ii ACKNOWLEDGEMENTS First of all, my thesis would not have been possible without the guidance and support from my supervisor Shawn Leroux. I also would like to express my gratitude to my co- supervisor and co-visionary Ken Oakes who first dreamed up this moose-stream project. The completion of this project is only possible due to the combined effort and encouragement from these two great research scientists. I thank Fred Baechler, Ian Fleming, and Matthew Smith, whose comments and suggestions greatly improved the body of work of this thesis. I also would like to thank Parks Canada for their funding and support, specifically I would like to thank Clayton D’Orsay and Rosie Smith for answering all my inquires about protocols and projects. A big thanks to James Bridgland for helping with data acquisition, as well as logistics and safety while trekking through the woodlands. Also, thanks to Michee Lemieux for GIS data acquisition and support of which greatly assisted with the progression of my project. I also would like to thank the great people at Nova Scotia Department of Natural Resources, namely Sean Basquill and Frances MacKinnon, as well as the Research Policy (RP) Grants Committee at Cape Breton University for continued funding for this project. Field work would not have been possible without the assistance of Caelin Murray. Thanks for having fun in the woods and streams with me, as well as saving my neck a few times. Additional thanks to Karla Boyd who assisted in the field when I first started this project, as well as Geoffrey MacEachen for technical support and assistance in the field. Big thanks to my dad, Irving MacSween, who let me borrow his ATV and truck, as well as helped in the field on occasion and who didn’t mind that I took over his garage for iii a summer. Thanks to my mother, Susanne MacSween for her support and encouragement again for my continued schooling. Finally, thank you to my lab mates: Jonathan Ebel, Semra Yalcin, Anne MacLeod, Matteo Rizzuto, and Sam Andrews, for the assistance, suggestions, and chocolate. iv TABLE OF CONTENTS ABSTRACT ........................................................................................................................ii ACKNOWLEDGEMENTS ...............................................................................................iii LIST OF FIGURES ..........................................................................................................vii LIST OF TABLES .............................................................................................................ix LIST OF APPENDICES ....................................................................................................xi Chapter 1: Introduction .....................................................................................................12 1.1. Herbivores as agents of disturbance ......................................................................12 1.2. Trophic interactions across ecosystem boundaries ................................................14 1.3. Impacts of forest removal on stream ecosystems ....................................................17 1.4. Thesis objectives, structure, and significance........................................................21 1.5. References .............................................................................................................23 Chapter 2: Watershed-scale effects of hyperabundant moose on stream ecosystems .......34 2.1. Introduction ...........................................................................................................34 2.2. Methods ..........................................................................................................................37 2.3. Results ............................................................................................................................46 2.4. Discussion ......................................................................................................................50 2.5. References ......................................................................................................................57 Chapter 3: Local-scale effects of hyperabundant moose on stream ecosystems ...............72 3.1. Introduction ....................................................................................................................72 3.2. Methods ..........................................................................................................................75 3.3. Results ............................................................................................................................86 v 3.4. Discussion ......................................................................................................................92 3.5. References ......................................................................................................................99 Chapter 4: Conclusions ...........................................................................................................124 4.1. Significance of Study ..................................................................................................124 4.2. Thesis Synopsis ...........................................................................................................124 4.3. Research Perspectives .................................................................................................128 4.4. Management Implications ..........................................................................................130 4.5. References ....................................................................................................................133 Appendix A: Chapter 2 Additional Data and Results ..........................................................138 Appendix B: Chapter 3 Additional Information, Data, and Results ..................................170 vi LIST OF FIGURES Figure 2.1: Watersheds included in the analysis for determining effects of grasslands on stream temperature in Cape Breton Highlands National Park ............................................64 Figure 2.2: Watersheds included in the analysis for determining effects of grasslands on total nitrogen, conductivity, and macroinvertebrates in Cape Breton Highlands National Park ............................................................................................................................................65 Figure 2.3: Relationships between a) Average maximum daily stream temperature, b) average stream temperature, and c) maximum stream temperature to percentage of grassland in 12 streams monitored during summer 2006-2016 .........................................66 Figure 2.4: Relationships between a) total nitrogen and b) electrical conductivity and percentage of grassland in 21 streams monitored during fall 2005-2014 ........................67 Figure 3.1: Headwater stream sites located in Cape Breton Highlands, Nova Scotia ..109 Figure 3.2: The locations of 7 grassland streams on North Mountain, Cape Breton Highlands National Park ......................................................................................................110 Figure 3.3: The locations of 7 forest streams located in various regions of the boreal forest of Cape Breton Highlands .........................................................................................111 Figure 3.4: Relationship between grassland (%) in the sub-basin and average daily temperature range for 13 headwater streams monitored July-September 2016 ............112 Figure 3.5: a) Total nitrogen in relation to grassland (%) for 14 headwater streams monitored monthly July-October 2016; b) Electrical conductivity in relation to grassland (%) for 12 headwater streams monitored monthly July-October 2016 (values overlap for forested sites) .....................................................................................................113 Figure 3.6: Chlorophyll a in relation to grassland (%) for 14 headwater
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