Castor Canadensis) Population Fluctuations, and Their Ecological Relationship with Salmonids
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Factors Influencing Beaver (Castor canadensis) Population Fluctuations, and Their Ecological Relationship with Salmonids A THESIS SUBMITTED TO THE FACULTY OF THE UNIVERSITY OF MINNESOTA BY Sean M. Johnson-Bice IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE Steve K. Windels, Adviser August 2019 © Sean M. Johnson-Bice 2019 ii Acknowledgments I wish to first thank my advisor and mentor Steve Windels for his continual guidance and patience, and the freedom he granted me to pursue my research interests. I would like to thank my committee members, George Host and Ron Moen, for their commitment and input in the progress of my thesis. Katti Renik and Andy Hafs both contributed significantly to the success of the first chapter, and Katti in particular assisted in writing initial portions of it. My second chapter simply would not have happened without the hard work and knowledge from Jake Ferguson, who conducted the statistical analysis, created two of the figures, and taught me all about Bayesian time series analysis in the process. I would also like to thank and acknowledge the contributions from Tom Gable and John Erb in my second chapter, who both provided tremendous insight and wisdom that greatly contributed to its development. My graduate school experience was a lot more enjoyable with the company of great friends, especially Ryan Steiner, Austin Homkes, Tom Gable, and Bradley Dawson. I also appreciate and acknowledge the financial support provided by the Minnesota Environment and Natural Resources Trust Fund, as recommended by the Legislative-Citizen Commission on Minnesota Resources (project M.L. 2016, Chp. 186, Sec. 2, Subd.03j); the University of Minnesota Duluth Integrated Biosciences Department Graduate Program summer fellowships awarded to me; and a one-semester teaching assistantship from the University of Minnesota Duluth Department of Biology. Part of my funding was also provided by the Coastal Zone Management Act of 1972, as amended, administered by the Office for Coastal Management, National Oceanic and Atmospheric Administration under Award NA18NOS190081 provided to the Minnesota Department of Natural Resources for iii Minnesota’s Lake Superior Coastal Program. Lastly, I want to thank several family members for their emotional support, including my brothers Alex and Brad Johnson- Bice; my aunt BJ Johnson and uncle Machan Shichi; my grandmother Sally Schofield; and of course my parents, David Bice and Kathleen Johnson, who have provided me with unwavering inspiration and encouragement. iv Dedication I dedicate this thesis to my biggest fans, my parents David Bice and Kathleen Johnson, who have always stood by side while I pursued my personal, professional, and above all, scientific endeavors. v Table of Contents List of tables ..................................................................................................................... viii List of figures ..................................................................................................................... ix Chapter 1: A Review of Beaver–Salmonid Relationships and History of Management Actions in the Western Great Lakes (USA) Region ................................................1 Summary ........................................................................................................................2 Introduction ....................................................................................................................3 History of Salmonids and Beavers in the Western Great Lakes Region .......................5 Review of Beaver Influence on Streams and Salmonids in the Western Great Lakes Region ....................................................................................................................13 Review of Beaver Management on Salmonid Streams in the Western Great Lakes Region ....................................................................................................................34 Management Implications ............................................................................................43 Conclusions ..................................................................................................................47 Tables ...........................................................................................................................51 Figures..........................................................................................................................53 Chapter 2: Factors Influencing Annual Rates of Change in the Number of Beaver Colonies ................................................................................................................55 Summary ......................................................................................................................56 Introduction ..................................................................................................................58 vi Methods........................................................................................................................62 Results ..........................................................................................................................72 Discussion ....................................................................................................................74 Management Implications ............................................................................................78 Tables ...........................................................................................................................80 Figures..........................................................................................................................83 Bibliography ......................................................................................................................87 vii List of Tables Table 1.1. Summary of the main effects found from 21 beaver-salmonid studies conducted within the WGL region .........................................................................53 Table 2.1. Summary of the 15 survey routes from northern Minnesota ............................80 Table 2.2. Comparison of three global models evaluating the influence of density- dependent and density-independent factors on growth rates of beaver colonies ...81 Table 2.3. Parameter estimates from the DDcov model ......................................................82 viii List of Figures Figure 1.1. Map showing where beaver–salmonid studies have been conducted in the western Great Lakes region ...................................................................................54 Figure 1.2. Timeline of major events from different management eras and a graph of the approximate beaver population trend from the western Great Lakes (WGL) region (1870–present) .......................................................................................................55 Figure 2.1. Map of the beaver survey study area and location of each route ....................83 Figure 2.2. Graphic depicting how survey routes were digitized by delineating the aircraft’s trajectory based on hand-drawn reference maps ....................................84 Figure 2.3. Composite image of raw (observed) data and model fits for each route ........85 Figure 2.4. Plot of the beta coefficients for the estimates from the full dataset compared to the estimates from the dataset with 1/3 of observations held out .....................86 ix CHAPTER ONE A Review of Beaver–Salmonid Relationships and History of Management Actions in the Western Great Lakes (USA) Region SUMMARY Within the western Great Lakes (WGL) U.S. region (Michigan, Minnesota, Wisconsin), the ecological impacts that North American beavers (Castor canadensis) have on cold- water streams are generally considered to negatively affect salmonid populations where the two taxa interact. Here, we review the history of beaver-salmonid interactions within the WGL region, describe how this relationship and management actions have evolved over the past century, and review all published studies from the region that have evaluated beaver-salmonid interactions. Our review suggests the impact beavers have varies spatially and temporally, depending on a variety of local ecological characteristics. We found beaver activity is often deleterious to salmonids in low-gradient stream basins, but generally beneficial in high- gradient basins; and ample groundwater inputs can offset the potential negative effects of beavers by stabilizing the hydrologic and thermal regimes within streams. However, there was an obvious lack of empirical data and/or experimental controls within the reviewed studies, which we suggest emphasizes the need for more data-driven beaver-salmonid research in the WGL region. Resource managers are routinely faced with an ecological dilemma between maintaining natural environmental processes within cold-water ecosystems and conducting beaver control for the benefit of salmonids, and this dilemma is further complicated when the salmonids in question are a non-native species. We anticipate future beaver-salmonid research will lead to a greater understanding of this ecologically-complex relationship that may better inform managers when and where beaver control is necessary to achieve the desired management objectives. 2 INTRODUCTION North American beaver (Castor canadensis) activities affect many fish and wildlife species (Rosell et al. 2005, Windels 2017), but of particular interest to resource managers in