SPATIAL ECOLOGY OF MOUNTAIN UNGULATES IN THE NORTHERN ROCKY MOUNTAINS: RANGE EXPANSION, HABITAT CHARACTERISTICS, NICHE OVERLAP, AND MIGRATORY DIVERSITY by Blake Henson Lowrey A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Fish and Wildlife Biology MONTANA STATE UNIVERSITY Bozeman, Montana November 2018 ©COPYRIGHT by Blake Henson Lowrey 2018 All Rights Reserved ii ACKNOWLEDGEMENTS This work was made possible through the dedicated support of many collaborators across state and federal agencies, universities, and NGOs. Bob Garrott served as a tireless mentor and inspiration through instilling his passions and expertise in ecology and wildlife management. P.J. White and Yellowstone National Park helped to secure the initial funding and facilitated capture and field logistics. The collaboration with Doug McWhirter and Wyoming Game and Fish greatly expanded the study scope, more than doubling our sample size throughout western WY. Alyson Courtemanch, Hollie Miyasaki, Kevin Monteith, and Sarah Dewey were instrumental in providing data from other regions throughout the Greater Yellowstone Area. Kelly Proffitt and Montana, Fish, Wildlife and Parks expanded our efforts throughout Montana. Lastly, countless biologists, wardens, and volunteers were instrumental to the project’s successes. Primary funding for this work was provided by the National Park Service, Canon USA Inc, the Wyoming Game and Fish Department, Federal Aid in Wildlife Restoration Grant W-159-R to Montana Fish Wildlife and Parks and the annual auction sale of a Montana bighorn sheep hunting license, Greater Yellowstone Coordinating Committee, the United States Forest Service, Montana State University, Wyoming Governor’s Big Game License Coalition, and Idaho Department of Fish and Game. Additional funds and scholarships were provided by Wyoming Wild Sheep Foundation, Montana Wild Sheep Foundation, Wild Sheep Foundation, International Order of Rocky Mountain Goats, Kevin Hurley Wild Sheep Biology Award, the Jack Creek Preserve Scholarship and Shikar Safari Club International. iii TABLE OF CONTENTS 1. INTRODUCTION TO THE DISSERTATION.............................................................. 1 2. SEASONAL RESOURCE SELECTION BY INTRODUCED MOUNTAIN GOATS IN THE SOUTHWEST GREATER YELLOWSTONE AREA ...................... 6 Contribution of Authors and Co-Authors ....................................................................... 6 Manuscript Information .................................................................................................. 7 Abstract ........................................................................................................................... 8 Key words ....................................................................................................................... 9 Introduction ..................................................................................................................... 9 Methods ........................................................................................................................ 12 Study area.................................................................................................................. 12 Animal capture and handling .................................................................................... 13 Data censoring .......................................................................................................... 13 Delineating mountain goat seasons ........................................................................... 15 Defining habitat availability ..................................................................................... 16 Resource covariates and spatial grain ....................................................................... 16 Statistical framework and model selection ............................................................... 19 Post-hoc model comparisons .................................................................................... 21 Mountain goat RSF extrapolations ........................................................................... 23 Model validation ....................................................................................................... 23 Results ........................................................................................................................... 24 Data collection, censoring, and definitions ............................................................... 24 Model selection and validation ................................................................................. 25 Mountain goat RSF extrapolations ........................................................................... 28 Discussion ..................................................................................................................... 29 Acknowledgements ....................................................................................................... 39 Tables and figures ......................................................................................................... 40 Appendix S2.1. Simulation methods to evaluate multiple sampling intensities when defining a used to available ratio for the study of resource selection by introduced mountain goats (Oreamnos americanus), southwest Greater Yellowstone Area, USA ............................................................................................... 45 Appendix S2.2. Details of individual mountain goats .................................................. 51 iv TABLE OF CONTENTS CONTINUED Appendix S2.3. Detailed results from the delineation of mountain goat seasons according to migration parameters estimated from nonlinear regression modeling of net-squared displacement ......................................................................................... 52 Appendix S2.4. Complete results from the tiered model selection process and post-hoc model evaluation ............................................................................................ 55 Appendix S2.5. Mountain goat GPS locations in relation to interpolated and extrapolated RSF predictions ........................................................................................ 66 Appendix S2.6. Seasonal k-fold cross-validation results for the Snake River Range study area, ID and WY, USA ............................................................................ 67 Appendix S2.7. Mountain goat RSF extrapolations to the southwest Greater Yellowstone Area ......................................................................................................... 69 3. NICHE SIMILARITIES AMONG INTRODUCED AND NATIVE MOUNTAIN UNGULATES ........................................................................................ 74 Contribution of Authors and Co-Authors ..................................................................... 74 Manuscript Information ................................................................................................ 75 Abstract ......................................................................................................................... 76 Introduction ................................................................................................................... 77 Methods ........................................................................................................................ 82 Study area.................................................................................................................. 82 Data collection and censoring ................................................................................... 83 Realized niche components....................................................................................... 84 Seasonal niche characterizations ............................................................................... 86 Evaluating niche similarity ....................................................................................... 88 Results ........................................................................................................................... 90 Data collection and censoring ................................................................................... 90 Seasonal niche characterization and similarity ......................................................... 90 Discussion ..................................................................................................................... 93 Acknowledgments ...................................................................................................... 100 Tables and figures ....................................................................................................... 101 Appendix S3.1: Delineating summer and winter seasons for sympatric bighorn sheep and mountain goats in the northeast Greater Yellowstone Area ...................... 108 v TABLE OF CONTENTS CONTINUED Appendix S3.2: Seasonal fix success for sympatric bighorn sheep and mountain goats in the northeast Greater Yellowstone Area ....................................................... 112 Appendix S3.3. Simulation methods to evaluate multiple sampling intensities when defining a used to available ratio for sympatric bighorn sheep and mountain goats in the northeast Greater Yellowstone Area ....................................... 114 Appendix S3.4: Individual summary statistics ........................................................... 118 Appendix