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INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6” x 9” black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. UMI A Bell & Howell Information Company 300 North Zed) Road, Ann Arbor MI 48106-1346 USA 313/761-4700 800/521-0600 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. FORAGING ECOLOGY OF FEMALE DALL’S SHEEP IN THE BROOKS RANGE, ALASKA A THESIS Presented to the Faculty of the University of Alaska in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY By Michael Charles Hansen, B.S., M.S. Fairbanks, Alaska May 1996 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. UMI Number: 9632222 UMI Microform 9632222 Copyright 19%, by UMI Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, MI 48103 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. FORAGING ECOLOGY OF FEMALE DALL’S SHEEP IN THE BROOKS RANGE, ALASKA by Michael Charles Hansen RECOMMENDED: Lo ■ fckt/C ttd O Advisory Committee Chair Vkk 1 . —-t, Department rHead \ APPROVED: Dean, College of Natural Resources Dean of the Graduate School Date Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ABSTRACT Most wild sheep (Ovis) are primarily diurnal. Thus, extreme cold, darkness, and limited quantities of low-quality forage during long winters above the Arctic Circle present a formidable challenge for sheep. Further, summer is particularly short at these high latitudes, providing little time for sheep to accumulate energy reserves for winter. This thesis discusses dietary and behavioral responses of wild sheep to the constraints of Arctic environments. Specifically, I determined diet composition and selection, forage quality, nutrient intake, and activity budgets of adult female Dali’s sheep (ewes) (Ovis dalli dalli) near the northern extreme of the range of wild sheep for 2 years and constructed a model of the energy relationships of these animals. Ewes primarily consumed forbs and grasses during summer, and strongly selected forbs over other forages in accordance with the predictions of optimal foraging theory. Diets primarily consisted of grasses in early winter, shifted to sedges in February, and back to grasses in early spring. Shrubs were consistently the least selected class of forage. When the diet was composed of forages with varying digestibility, microhistologjcal analyses not corrected for differential digestibility were biased toward less digestible forage. Winter forage available to Dali’s sheep in the northern Brooks Range was low in both digestibility and protein content. In early summer ewes foraged during all hours of the day when sunlight was present for 24 hours. Sheep restricted their foraging almost entirely to daylight hours near the equinoxes, and foraged during all available hours of light, as well as 2.8 hours of the Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. night in December. Daily foraging time varied from 12.9 hours in June to 7.9 hours in December, and, when measured on a daily basis, was positively correlated with average windchill and daylength. Ash-free fecal nitrogen and in vitro digestible dry matter were most highly correlated with activity level on a monthly basis. Energetics modeling indicated that ewes were in a negative energy balance for 6-8 months each winter and lost nearly 30% of their body weight. Duration of the short summer growing period was most important for weight gain, and presence of deep snow determined weight loss in winter. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. TABLE OF CONTENTS ABSTRACT....................................................................................................................... iii TABLE OF CONTENTS................................................................................................... v LIST OF FIGURES............................................................................................................ix LIST OF TABLES............................................................................................................xii INTRODUCTION...............................................................................................................1 CHAPTER 1. COMPARISON OF THREE METHODS FOR EVALUATING ACTIVITY OF DALL’S SHEEP............................................................................6 ABSTRACT.................................................................................................................... 6 INTRODUCTION...........................................................................................................7 STUDY AREA................................................................................................................8 METHODS..................................................................................................................... 9 Data Collection ............................................................................................................9 Analyses......................................................................................................................11 Calibration ............................................................................................................. 11 Validation .............................................................................................................. 12 Cost Comparison .......................................................................................................13 RESULTS......................................................................................................................14 VHF Telemetry vs. Direct Observation ...................................................................... 14 Calibration ............................................................................................................. 14 Validation .............................................................................................................. 15 Satellite Telemetry vs. Direct Observation ................................................................. 18 Calibration ............................................................................................................. 18 Validation ..............................................................................................................20 Cost Comparison ...................................................................................................... 20 DISCUSSION...............................................................................................................21 VHF Telemetry..........................................................................................................21 Satellite Telemetry.................................................................................................... 23 MANAGEMENT IMPLICATIONS.............................................................................24 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LITERATURE CITED 25 CHAPTER 2. DAILY FECAL EXCRETION AND DEFECATION RATES OF FREE-RANGING DALL’S SHEEP CALIBRATED USING DOMESTIC SHEEP................................................................................................................ 29 ABSTRACT................................................................................................................ 29 INTRODUCTION.......................................................................................................30 STUDY AREA............................................................................................................33 METHODS................................................................................................................. 33 Domestic Sheep .......................................................................................................33 Dali’s