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RISK EFFECTS in ELK (Cervus Elaphus): BEHAVIORAL and NUTRITIONAL

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RISK EFFECTS in ELK (Cervus Elaphus): BEHAVIORAL and NUTRITIONAL RISK EFFECTS IN ELK (Cervus elaphus): BEHAVIORAL AND NUTRITIONAL RESPONSES TO WOLVES AND ENVIRONMENTAL CONDITIONS by David Alan Christianson A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Ecology and Environmental Sciences MONTANA STATE UNIVERSITY Bozeman, Montana April 2008 ©COPYRIGHT by David Alan Christianson 2008 All Rights Reserved ii APPROVAL of a dissertation submitted by David Alan Christianson This dissertation has been read by each member of the dissertation committee and has been found to be satisfactory regarding content, English usage, format, citation, bibliographic style, and consistency, and is ready for submission to the Division of Graduate Education. Dr. Scott Creel Approved for the Department of Ecology Dr. Dave Roberts Approved for the Division of Graduate Education Dr. Carl A. Fox iii STATEMENT OF PERMISSION TO USE In presenting this dissertation in partial fulfillment of the requirements for a doctoral degree at Montana State University, I agree that the Library shall make it available to borrowers under rules of the Library. I further agree that copying of this dissertation is allowable only for scholarly purposes, consistent with “fair use” as prescribed in the U.S. Copyright Law. Requests for extensive copying or reproduction of this dissertation should be referred to ProQuest Information and Learning, 300 North Zeeb Road, Ann Arbor, Michigan 48106, to whom I have granted “the exclusive right to reproduce and distribute my dissertation in and from microform along with the non- exclusive right to reproduce and distribute my abstract in any format in whole or in part.” David Alan Christianson April 2008 iv TABLE OF CONTENTS 1. INTRODUCTION .........................................................................................................1 2. A REVIEW OF ENVIRONMENTAL FACTORS AFFECTING ELK WINTER DIETS............................................................................................................6 Abstract..........................................................................................................................6 Introduction....................................................................................................................7 Winter Range Habitat Type .......................................................................10 Severity of Winter Conditions ...................................................................10 Methods of Determining Diets...................................................................11 Predation Risk (Human Hunting) ..............................................................12 Period of Winter.........................................................................................12 Methods........................................................................................................................13 Dependent Variable - Diet Composition....................................................14 Independent Variables ...............................................................................16 Winter Range Habitat Type ...........................................................16 Severity of Winter Conditions .......................................................17 Methods of Determining Diets.......................................................17 Predation Risk (Human Hunting) ..................................................17 Period of Winter.............................................................................18 Statistical Analysis.....................................................................................18 Results..........................................................................................................................19 Broad Diet Patterns and Comparison with Sympatric Ruminants.............19 Effects of Range Type and Winter Severity ..............................................23 Effects of Study Methods, Hunting, and Period of Winter........................26 Methods of Determining Diets.......................................................26 Predation Risk (Human Hunting) ..................................................27 Period of Winter.............................................................................28 Interactions.....................................................................................28 Discussion....................................................................................................................29 Management Implications............................................................................................33 Acknowledgements......................................................................................................34 3. THE OPTIMAL BALANCE BETWEEN GRAZING AND BROWSING IN ELK FROM MODELS OF MASS DYNAMICS........................................................35 Abstract........................................................................................................................35 Introduction..................................................................................................................36 Methods........................................................................................................................42 Raw Data on Body Mass Changes Under Experimental Diets..................42 Diet Composition...........................................................................43 Metabolic Mass..............................................................................43 v TABLE OF CONTENTS continued Dietary Nitrogen ............................................................................44 Feeding Situation ...........................................................................44 Intake Rate .....................................................................................44 Model Selection .........................................................................................45 Results..........................................................................................................................48 Experimental Diets.....................................................................................48 Selection Among Regression Models for Body Mass Change..................50 Predicting the Optimal Balance of Grazing and Browsing........................52 Discussion....................................................................................................................56 4. FECAL CHLOROPHYLL REVEALS COMPLEX RELATIONSHIPS BETWEEN PRIMARY PRODUCTION AND CONSUMPTION IN A TEMPERATE HERBIVORE ......................................................................................64 Abstract........................................................................................................................64 Introduction..................................................................................................................65 Methods........................................................................................................................70 Study Area and Data Collection ................................................................70 Fecal and Forage Samples .............................................................71 Sample Extraction and Nutritional Analysis: ................................72 Snow and Temperature ..................................................................73 Primary Production ........................................................................73 Spectrophotometric Technique for Assay of Chlorophyll in Fecal and Grass Extracts................................................................................75 Impetus and Validation..................................................................75 Primary Measurement of Chlorophy .............................................77 Data Analysis.............................................................................................78 Describing Temporal Patterns........................................................78 Describing the Relationship Between NDVI and Fecal Chlorophyll ..............................................................................80 Statistical Analysis.........................................................................81 Results..........................................................................................................................82 Plant and Fecal Light Absorption ..............................................................82 Temporal Trend in Winter Conditions and Fecal Chlorophyll..................84 Comparing Fecal Chlorophyll With Temporal Trends in Fecal Nitrogen ...............................................................................................88 Comparing Fecal Chlorophyll With Trends In Grass Nitrogen, Chlorophyll, and Bite Size...................................................................90 Comparing Fecal Chlorophyll with Temporal Trends in Primary Productivity..........................................................................................92 Relationship Between Primary Productivity and Fecal Chlorophyll.........94 Discussion....................................................................................................................98 vi TABLE OF CONTENTS continued 5. RISK EFFECTS IN ELK: SEX-SPECIFC RESPONSES IN GRAZING AND BROWSING DUE TO PREDATION RISK FROM WOLVES...............................103 Abstract......................................................................................................................103
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