Winter Ecology of Brown Trout in Groundwater Buffered Streams a DISSERTATION SUBMITTED to the FACUL
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Protected from the Elements: Winter Ecology of Brown Trout in Groundwater Buffered Streams A DISSERTATION SUBMITTED TO THE FACULTY OF THE UNIVERSITY OF MINNESOTA BY William E French IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY Bruce Vondracek and Leonard C. Ferrington Jr., advisers September, 2014 © William E. French, 2014 Acknowledgements I would like to take a moment to thank the people and organizations which have made this dissertation possible. Your continued assistance and support have been critical to the pursuit of my educational goals. First, I would like to thank my co- advisors, Bruce Vondracek and Len Ferrington Jr. Len has been a valuable asset to my dissertation research, especially during the planning and study design phases. Len’s expertise with aquatic invertebrates was a great help during the identification of trout stomach contents. His knowledge and enthusiasm regarding aquatic insects have inspired me to think more holistically about aquatic ecosystems: invertebrates are much more than simply fish food. Bruce has had an important influence on my professional development since I was an undergraduate, his door has always been open. He was always willing to review a potential publication or encourage me to attend a meeting to present my research. Bruce was always willing to pitch in and help out in the field, even in the dead of winter. He made a point to get out sampling with us on at least one trip each winter. I have always admired the passion he has for fisheries science, and hope to emulate this in my own career. Whether our discussions focused on a change in sampling protocol, a recent fishing trip, or Monday morning quarterbacking the latest Packer or Gopher Hockey game; a conversation with Bruce was always enjoyable. I am proud to say I consider him a friend as well as an advisor and mentor, and will miss the time I was able to spend with him. Thanks also to the rest of my committee, Doug Dieterman and Jacques Finlay for their guidance and support during the past four years. Jacques was a great help with i the stable isotope component of my dissertation, and helped with planning, analyses, and discussions of results. Doug was very involved with selection of sampling sites and his knowledge of Brown Trout ecology was a great help in guiding the direction of my research. Doug and I shared many conversations regarding what I was seeing in my work and how it could relate to trout ecology and management in southeastern Minnesota. I would like to extend a special thanks to my fellow graduate students and undergraduate volunteers who were part of our winter field work team, especially Jane Mazack and Patrick Sherman. We shared many long cold days together (and kept the Creamery and the Sawmill Inn in business through the winter) during our three winters in southeastern Minnesota, and I thoroughly enjoyed our field time. Your camaraderie played a large part in that. I would like to thank the MN DNR staff of the Lanesboro and Lake City offices, especially John Hoxmeier and Dan Spence for their assistance, support, and advice throughout this project. Minnesota DNR provided guidance on stream trout populations and site access, loaner equipment when some had been forgotten, and often helped us out as an extra pair of hands in the field. Thanks to the faculty and staff in the Conservation Biology Graduate Program and the US Geological Survey Cooperative Fish and Wildlife Research Unit, the Water Resources Center, and the Graduate School at the University of Minnesota for their help and assistance throughout the past four years. A special thanks to Rob Blair and Karen ii Oberhauser for their efforts helping me navigate the logistical side of the Graduate School during their time as DGS. Thank you to Hattie Saloka, David Fulton and David Anderson for their support and friendship. I did my best to keep the candy on Hattie’s desk fresh, and compare our latest notes on the Grouse season (or the turkey, duck, pheasant, deer, or rail season) with David Anderson and David Fulton. I especially want to thank my friends, office mates, and fellow cohort members in the Conservation Biology program. I have thoroughly enjoyed the past four years and you all have been a big part of that. Cons Bio parties and potlucks, happy hours, random nights out on the town have all been memorable. I particularly enjoyed our unofficial ice fishing club and a special thanks to those of you who I’ve wet a line with over the past couple years. Finally, I want to thank my parents, extended family, and especially my wife Jennifer for her continued support and encouragement as I have pursued my educational goals. She has constantly stood by me, waiting patiently for her husband to finally grow up, graduate and get a real job. Thank you again to you all. I would not have been able to get this far without your continued love and support. iii Abstract Winter has traditionally been considered a period of dormancy for stream dwelling trout in temperate latitudes. Seasonal changes including low water temperatures, ice formation, and reduced prey availability from aquatic and terrestrial sources often contribute to reductions in trout growth and survival. Consequentially, winter has rarely been the focus of study by fisheries scientists, and relatively little information is available regarding stream trout during winter. However, because of the potential impact on stream trout growth and survival, winter is an important season for trout populations and of particular interest to fisheries managers. The goal of this dissertation was to examine winter trophic ecology of stream dwelling trout populations, and the potential of groundwater input to buffer stream water temperatures and trout from the environmental conditions typically associated with winter. This dissertation consists of three chapters that contribute towards this goal. The first chapter examines winter diet of Brown Trout by quantifying trout consumption, identifies important winter prey taxa , and compares diet composition among a number of trout populations. The second chapter describes Brown Trout winter growth and condition, and examines the influence of groundwater buffering and trout diet composition on growth and condition. The third and final chapter uses stable isotope analyses to examine seasonal variation in Brown Trout diets, and the position of trout within winter food webs of groundwater dominated streams. My dissertation will help managers predict the potential effects of winter on important recreational stream iv trout fisheries, and allow informed management decisions incorporating the best available information. v Contents Acknowledgements i Abstract iv List of Tables vi List of Figures vii Prologue 1 Chapter 1: Winter diet of Brown Trout Salmo trutta in groundwater-dominated streams of southeastern Minnesot 7 Introduction .............................................................................................................................................. 9 Materials and Methods ........................................................................................................................... 12 Results ..................................................................................................................................................... 16 Discussion ............................................................................................................................................... 19 Chapter 2: Winter growth and condition of Brown Trout Salmo trutta in groundwater- dominated streams 38 Introduction ............................................................................................................................................ 40 Methods .................................................................................................................................................. 44 Results ..................................................................................................................................................... 47 Discussion ............................................................................................................................................... 48 Conclusions ............................................................................................................................................. 51 Chapter 3: Use of stable isotope analyses to examine seasonal trophic relationships of Brown Trout in groundwater dominated streams 61 Introduction ............................................................................................................................................ 63 Methods .................................................................................................................................................. 67 Results ..................................................................................................................................................... 69 Discussion ............................................................................................................................................... 71 Bibliography 91 vi List of Tables 1.1 Sampling information and streams included in Brown Trout winter diet analyses….....26 1.2a Prey taxa by number from sampled Brown Trout during early winter………………….……..27 1.2b Prey taxa by number from sampled Brown Trout during late winter……………………………29 1.3a Prey taxa by dry weight from sampled Brown Trout during early winter……………………..31 1.3b Prey