Winter ecology of moose on the Northern Yellowstone Winter Range by Daniel Bruce Tyers A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Fish and Wildlife Biology Montana State University © Copyright by Daniel Bruce Tyers (2003) Abstract: Shiras moose (Alces alces shirasi) invaded the Northern Yellowstone Winter Range (NYWR) in the 1800’s. Management of this species has been handicapped by limited population data and reliance on habitat use paradigms from other areas in North America. For example, although moose in most regions forage in recently disturbed forests with abundant deciduous shrubs, I did not find this to be true on the NYWR. I used multiple methods to determine population status, including aerial surveys, horseback surveys, road surveys, and ground counts in willow habitats. Moose were most easily seen when they foraged in willow (Salix spp.) stands in November-December and May-June. Attempts to count moose at other times resulted in marked under-sampling. Fires in 1988 removed mature forests at a landscape level resulting in a loss of preferred winter habitat and a sustained population reduction. I also investigated moose winter foraging activities among burned (1988), logged, forested, and forest edge environments. Logged areas and recent bums were rarely used, but forest interiors were important habitat. The edge effect did not benefit moose in winter. NYWR moose coped with winter by seeking concentrations of food to maximize feeding and minimize movement. They optimized foraging by using 2 taxa that grow in patches, willow and subalpine fir (Abies lasiocarpa). Moose fed in willow until snow forced them to nearby forested hillsides where they browsed subalpine fir <5 m in height. Subalpine fir, a shade tolerant tree, grows more rapidly under a forest canopy. It can reproduce through layering, creating an expanding patch of small trees around the parent. Patches are largest and most numerous in the oldest lodgepole pine (Pinus contorta) forests. Moose used other characteristics of lodgepole forests to reduce energy output by finding routes with less than average snow depth to reach subalpine fir patches. Moose also browsed gooseberry (Ribes spp.) and buffaloberry (Shepherdia canadensis), nutrient rich shrubs that are scattered and energetically expensive to obtain. Exclosure studies showed willow was suppressed before 1988, and fires and drought increased negative effects. Higher willow utilization rates in 1989 probably reflected a response by moose to a loss of winter habitat due to the 1988 fires, a foraging strategy that may have contributed to observed willow mortality. From 1989 to 1997, some willow showed signs of recovery. Replacement of older forests where moose foraged before 1988 could take several hundred years. Maintaining remaining populations should concentrate on minimizing loss of mature forests and adjacent willow and careful hunting management.  WINTER ECOLOGY OF MOOSE ON THE NORTHERN YELLOWSTONE WINTER RANGE by Daniel Bmce Tyers 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 April 2003 D3'75> T cUfeI ii APPROVAL of a dissertation submitted by Daniel Tyers This dissertation has been read by each member of the dissertation committee and has been found to be satisfactory regarding content, English usage, format, citations, bibliographic style, and consistency, and is ready for submission to the College of Graduate Studies. Jay Rotella Approved for the Department of Ecology (Signature) Approved for the College of Graduate Studies Bmce McLei iignature) Date 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 Bell & Howell 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.” Signature Date ACKNOWLEDGMENTS iv I would like to thank the National Park Service, U.S. Forest Service, Montana Fish, WildUfe and Parks, John and Kathryn Harris, Ingrid Vimont, and Safari Club International for financial support. Tom Erskine represented Safari Club, Montana Chapter. I would also like to thank the members of my committee: Lynn Irby, Tom McMahon, Harold Picton, Jack Taylor, and Peter Gogan. Many National Park Service employees participated or endorsed this project including: John Varley, Frank Singer, Stu Coleman, Dan Sholly, Sue Console Murphy, Tom Olliff, Jennifer Whipple, Laurel Mack, Barry Jordan, Randy and Sally King, Andy Fisher, Mike Phlaum, John Donaldson, Bill Goe, Brian O’Dea, Doug Smith, Gary Brown, Roy Renkin, and Craig McClure. I am indebted to Montana Fish, Wildlife, and Parks employees Ainie Foss, Kurt Alt, Tom Lemke, Joel Petersen, Glenn Erickson, Hany Whitney, Mike Ross, and Ken Hamlin. I would credit Tom Puchlerz, U.S. Forest Service, for being the creative mind and primary catalyst for this study. Other Forest Service employees who contributed include: Marion Cherry, Mark Petroni, JT Stangl, Patrick Hoppe, Rich Inman, Bob Gibson, Larry Lewis, and Colleen Stein. These were the troops in the field or behind the computer: Sam Reid, Steve Yonge, Par Hermansson, Jeremy Zimmer, Nicolle Smith, Sarah Cobum, Chief, Molly, Reyer Rens, Kim Olson, Jeannie Heltzel, Maria Newcomb, Johan Kers, Cynthia Collin, TomNelson, Rob St. John, Vanna Boccadori, Melanie Weeks, Matthew Tyers, Debby Young, Jack Hopkins, Kathy McMahon, Phyllis Birdwell, Maiy Beth Gujda, Rachael Ondov, Mike Beebe, Anna-CarinAndersson, and Emma Cayer. Phyllis Wolfe and Tami Blackford made invaluable contributions to several data sets. Bill Gasaway, Jim Peek, Dave Mattson, Bonnie Blanchard, and Sandi Fowler gave important advice. I reUed on Dave Stringfield, Tom Drummer, and Scot Shuler for statistical help. Bill Chapman, SagebrushAero, flew telemetry and survey flights. Laszlo Torma and the staff at the MSU analytical laboratory conducted nutritional analysis on food samples. Drs. Jim Murray, Victor Van Ballenberghe, and Tom Thome gave advice with animal immobilization. I would like to thank the Silver Tip Ranch for then help, especially Shirley Blakeley and Steve Garvey. Warren Johnson and Dave Poncin kept track of backcountry moose population trends. This project could not have been completed without the remarkable dedication of Tris Hoffman and Ellen Snoeyenbos. In memory of Jodi Moore, Abrey Haines, Bill Gasaway, Larry Lewis, Scotty Chapman, and Amie Foss. I wish they could be here to see the final product. My profound apologies and regrets to family and friends who have been put on hold for the past 18 years. V TABLE OF CONTENTS 1. LIST OF TABLES................................................................................................................................... ... 2. LIST OF FIGURES..............................................................................................................................xxv 3. ABSTRACT................................................. :.................................................................................... xxxi 4. INTRODUCTION.................................................................................................................................... I Project Origin...........................................................................................................................................I Background Information......................................................................................................................... 2 Project Organization.............................................................................................................................. 8 5. STUDY AREA......................................................................................................................................... 9 Soda Butte Study Unit......................................................................................................................... 10 Slough Creek Study Unit..................................................................................................................... 12 Bear Creek Study Unit......................................................................................................................... 13 Yellowstone Park Study Un it ............................................................................................................. 14 6. MOOSE POPULATION DESCRIPTION AND MONITORING ON THE NORTHERN YELLOWSTONE WINTER RANGE................................................................................................... 15 Introduction.......................................................................................................................................... 15 Methods............................................................................................................................................. 16 Historical Documents........................................................................................................................ 16 Horseback Transect Index.................................................................................................................. 16 Road Transect Index.......................................................................................................................... 16 Willow Stand Over-flight Index........................................................................................................ 18 Daily Willow Stand Observations..................................................................................................... 18 Census Flights.................................................................................................................................... 19 Statistical Tests.................................................................................................................................. 19 Results.................................................................................................................................................... 20 Historical Documents............................................... 20 Horseback Transect Index..................................................................................................................23 Road Transect Index.......................................... 24 Willow Stand Over-flight Index........................................................................................................ 27 Daily Willow Stand Observations......................................... 28 Census Flights.................................................................................................................................... 30 Discussion.............................................................................................................................................. 31 Population History..... :...................................................................................................................... 31 Optimal Population Monitoring.........................................................................................................32 7. MOOSE HOME RANGE CHARACTERISTICS AND SURVIVAL ON THE NORTHERN YELLOWSTONE WINTER RANGE................................................................................................... 37 Introduction.................... ......................................:...................... ;..................................................... 37 Methods................................................................................................................................................. 41 Animal Capture and Radio Locations................................................................................................ 41 Home Range Analysis........................................................................................................................ 41 Survival of Radiocollared Moose......................................................................................................42 Statistical Tests.................................... 43 Results................................................................................................................................................... 43 Home Range Size.............................................................................................................................. 43 Home Range Configmation............................................................................................................... 44 Habitat Changes................................................................................................................................. 45 Survival of Radiocollared Moose................................................. i.................................................. 47 Discussion..................................................................................................................... 48 Home Range Size.......................................................................... 48 Survival of Radiocollared Moose...................................................................................................... 50 Ecological Implications..................................................................................................................... 50 8. MOOSE DIET AND FORAGING PATTERNS ON THE NORTHERN YELLOWSTONE WINTER RANGE................................................................................................................................. 53 Introduction.......................................................................................................................................... 53 Methods............................................................................................................................................. 54 Browse Use Variation........................................................................................................................ 56 Use of Browse in Relationship to Abundance................................................................................... 56 Variation in Browse Use in Cover Categories and Among Successional Stages............................. 56 Variation in Browsing Rate by Cover Category................................................................................ 57 Variation in Browsing on Subalpine F ir ............................................................................................ 57 Nutritional Analysis of Browse Species............................................................................................ 57 Statistical Tests.................................................................................................................................. 58 Results................................................................................................................................................... 58 Diet Composition Variation............................................................................................................... 58 Use of Browse in Relationship to Abundance................................................................................... 62 Variation in Browsing Among Cover Categories.............................................................................. 62 Variation in Browsing Rate in Cover Categories............................................................................... 66 Variation in Use of Subalpine F ir ........................ 67 Nutritional Analysis of Browse Species............................................................................................ 69 Discussion.............................................................................................................................................. 74 9. MOOSE COVER TYPE USE ON THE NORTHERN YELLOWSTONE WINTER RANGE........... 79 Introduction.......................................................................................................................................... 79 Methods.......................................................................................................................................... 81 Variation in Cover Category Use by Radiocollared Moose............................................................... 82 Location of Pellet Groups and Moose Beds by Cover Category..................................................... 82 Variation in Cover Category Use by Moose Seen Along the Cooke City Road...............................83 Variation in Cover Categoiy Selection of Moose and Sympatric Cervids on Track-Intercept Transects.......................... S3 Statistical Tests.................................................................................................................................. 85 Results................................................................................................................................................... 86 Variation in Cover Category Use by Radiocollared Moose Among Winter Periods......................... 86 Variation in Cover Category Use by Radiocollared Moose Among Study Units.............................. 87 Variation in Cover Category Use by Radiocollared Moose Related to Fire Effects.......................... 88 Distribution of Moose Pellet Groups and Beds Among Cover Categories....................................... 94 Variation in Cover Category Use by Moose SeenAlong the Cooke City Road...............................95 Moose Cover Type Selection on Track-Intercept Transects.............................................................. 99 Elk Cover Type Selection on Track-Intercept Transects................................................................. 102 Mule Deer Cover Type Selection oh Track-Intercept Transects..................................................... 104 Discussion.............................................................................................................................................104 Variation in Moose Cover Category Use......................................................................................... 104 Variation in Cover Category Use Among Sympatric Cervids on Moose Winter Range................ 109 vi 10. THE EFFECTS OF SNOW ON MOOSE HABITAT USE ON THE NORTHERN YELLOWSTONE WINTER RANGE................................................................................................ I l l Introduction........................................................................................................................................I l l Methods.................................................................................................................................................113 Variation in Snow Depth at Moose Beds and Travel Routes Among Winter Periods....................113 Average Height of Browsed Shrubs Compared to Average Snow Depth.......................................114 Variation in Snow Depth by Moose and Sympatric Cervids on Track-Intercept Transects........... 114 Results..................................................................................................................................................116 Variation in Snow Depth at Moose Beds and Travel Routes Among Winter Periods...................116 Average Height of Browsed Shrubs Compared to Average Snow Depth.......................................117 Variation in Snow Depth Selection by Moose on Track-Intercept Transects.................................117 Variation in Snow Depth Use by Elk on Track-Intercept Transects.................................................119 Variation in Snow Depth Use by Mule Deer on Track-Intercept Transects...................................121 Discussion............................................................................................................................................ 122 11. FACTORS INFLUENCING THE DISTRIBUTION OF MOOSE IN CONIFER FORESTS ON THE NORTHERN YELLOWSTONE WINTER RANGE........................................................... 126 Introduction.........................................................................................................................................126 Methods................................................................................................................................................ 127 Tree and Shrub Characteristics of NYWR Moose Feeding Sites in Forests....................................127 Ungulate Activity and Subalpine Fir Characteristics in Representative Lodgepole Pine Forests of Different Ages................................................................................................................. 128 Statistical Tests.................................................................................................................................130 Results..................................................................................................................................................130 Tree and Shrub Characteristics of Moose Feeding Sites in Forests................................................ 130 Ungulate Activity and Subalpine Fir Characteristics in Representative Lodgepole Pine Forests of Different Ages................................................................................................................. 136 Discussion.............................................................................................................................................138 Forage Characteristics of Forested Moose Winter Range............................................................... 138 Foraging Theoiy and NYWR Moose Winter Distribution in Conifer Forests................................ 140 12. IMPACTS OF CLEARCUTS ON MOOSE HABITAT USE IN LODGEPOLE PENE FORESTS ON THE NORTHERN YELLOWSTONE WINTER RANGE........................................................... 143 Introduction.........................................................................................................................................143 Methods................................................................................................................................................145 Statistical Tests................................................................................................................................ 147 Results..................................................................................................................................................147 Pellet Group Distribution................................................................................................................. 147 Shrub Characteristics........................................................................................................................149 Tree Characteristics...................................................................... :................................................ 152 Discussion............................................................................................................................................ 159 13. WILLOW AVAILABILITY AND WILLOW AS A DETERMINANT OF MOOSE HABITAT USE ON THE NORTHERN YELLOWSTONE WINTER RANGE.................................................. 164 Introduction........................................................................................................................................ 164 Methods............................................................................................................................................... 166 Characteristics of Moose Feeding Sites With Willow..................................................................... 166 Monthly Variation in Snow Conditions and Ungulate Activity in Willow Stands.......................... 167 Variation in Willow Survival, Height, Percent Browsing, and Twig Production............................ 168 Variation in Willow Height and Percent Cover In and Out of the Slough Creek Exclosure............169 Statistical Tests............................................. 169 vii Viii R e s u l t s .....................................................................................................................................................................................................170 C h a ra c te r is t ic s o f M o o s e F e e d in g S ite s W ith W i l l o w ............................................................................................ 170 M o n th ly V a r ia t io n i n S n o w C o n d itio n s a n d U n g u la te A c t iv i ty in 8 W illo w S t a n d s ..............................171 W illo w H e ig h t V a r ia tio n i n 7 S t a n d s ................................................................................................................................. 173 W illo w B ro w s in g R a te V a r ia tio n in 7 S t a n d s ............................................................................................................. 175 W illo w T w ig P ro d u c t io n V a r ia tio n in 7 S t a n d s ......................................................................................................... 178 W illo w S u rv iv a l V a r ia tio n i n 7 S ta n d s .............................................................................................................................. 181 V a r ia tio n i n W illo w S ta n d F ire E ffe c ts A m o n g S tu d y U n i t s ................................................................... ..........182 V a r ia tio n in W il lo w H e ig h t a n d P e rc e n t C o v e r I n a n d O u t o f th e S lo u g h C re e k E x c lo s u r e ..............183 D is c u s s io n ..............................................................................................................................................................................................185 14. C O N C L U S I O N ........................... 190 M o o se P opulation S t a t u s ..............................................................................................................................................................190 M o o se H abitat S ta tu s ......................................................................................................................................................................193 M o o se W in ter Strategy in th e N Y W R ................................................................................................................................. 198 M o o se H abitat M a n a g e m e n t ...................................................................................................................................................... 201 S u m m a r y ................................................................................................................................................................................................ 2 0 4 15. L IT E R A T U R E C I T E D .......................................................................................................................................................... 2 0 6 A P P E N D I C E S ............................................................................................................................................................................................. 2 2 3 A P P E N D IX A : post hoc te st d eterm in in g sign ifican t d ifferences in t h e average n u m ber OF MOOSE SEEN PER 1 /2 HOUR OVER A 6 4 WEEK PERIOD IN A SODA BUTTE WILLOW STAND .................................2 2 4 A P P E N D IX B : n u m b er o f m o o se seen on survey flig hts o n th e northern Yello w sto ne w in ter ra n g e fr o m 19 6 8 - 1 9 9 2 ..........................................................................................................................................2 2 6 A P P E N D IX C : post hoc test d eterm in in g sign ifican t differences in th e average m oo se h o m e r a n g e size AMONG 4 STUDY UNITS............................................................................................................................... 2 2 8 A P P E N D IX D : pla n t species a ppea rin g in th e d ie t o f m o o s e .................................................................................. 2 3 0 A P P E N D IX E : pla n t species ea ten b y m oo se ex pressed a s a percentage of th e total d iet DURING THE EARLY, M B -, AND LATE WINTER PERIODS .......................................................................... 2 3 2 A P P E N D IX F : pla n t species eaten b y m oo se ex pressed a s a percentage of th e total d iet PRE- AND POST-FIRE DURING THE EARLY, M B -, AND LATE WINTER PERIODS...................................:........................... 2 3 4 A P P E N D IX G : pla n t species eaten b y m o o se ex pressed as a percentage o f th e total d iet PRE-AND POST-FIRE IN 4 STUDY UNITS .................................................................................................................................... 2 3 6 A P P E N D IX H : chi- squ a re a nalysis determ in in g if 5 tree species a re bro w sed b y m oo se IN PROPORTION TO AVAILABILITY IN SUCCESSIONAL STAGES OF 3 FORESTS TYPES .....................................................2 3 8 A P P E N D IX I: d istribu tio n of m o o se bro w sing a m o n g co ver categories d u r in g th e early , M B -, AND LATE WINTER PERIODS ............................................................................................................................................... 2 4 2 A P P E N D IX J: d istribution of m o o se bro w sing a m o n g co ver categories pr e- a n d po st-fire DURING THE EARLY, M B -, AND LATE WINTER PERIODS .......................................................................................................2 4 4 A P P E N D IX K : d istribu tio n of m o o se bro w sing a m o n g co ver categories pr e- a n d po st-fire IN 4 STUDY UNITS............................................................................................................................................................................. 2 4 6 A P P E N D IX L : post hoc test d eterm in in g significant differences in th e a verage n u m b er of TWIGS BROWSED PER SUBALPINE FIR AMONG WINTER PERIODS AND STUDY UNITS AND PRE- AND POST-FIRE ... 2 4 8 A P P E N D IX M : proxim ate a n d elem en ta l n utritio na l analysis of 5 m o o se bro w se s pe c ie s ..................2 5 0 A P P E N D IX N : ch i- squ a re analysis d eterm in in g if rado ico lla red m o o se in th e stu dy area WERE FOUND DURING THE WINTER IN COVER CATEGORIES PRE- AND POST-FIRE IN PROPORTION TO THEIR AVAILABILITY......................................................................................................................................................................... 2 5 3 A P P E N D IX O : chi- square analysis d eterm in in g if rado ico lla red m oo se w ere fo u n d during THE WINTER IN COVER CATEGORIES PRE- AND POST-FIRE AMONG 4 STUDY UNITS IN PROPORTION TO THEIR AVAILABILnY 255 ix APPENDIX P: c h i- s q u a r e a n a l y sis d e t e r m in in g if r a d o ic o l l a r e d m o o s e in 4 s t u d y u n it s WERE FOUND DURING THE WINTER IN AREAS WITH DISPARATE FIRE EFFECTS IN PROPORTION TO THEIR AVAILABILITY.............................................................................................................................................................................................. 258 APPENDIX Q: c h i- s q u a r e a n a l y sis d e t e r m in in g if m o o s e , e l k , a n d m u l e d e e r t r a c k s a p p e a r DURING THE WINTER IN COVER CATEGORIES ALONG FIXED TRANSECTS IN PROPORTION TO THEIR AVAILABILITY..................................................................................................................................................................................260 APPENDIX R: c h i- s q u a r e a n a l y s is d e t e r m in in g if m o o s e , e l k , a n d m u l e d e e r t r a c k s a p p e a r IN SNOW DEPTH CATEGORIES ALONG FIXED TRANSECT IN PROPORTION TO THEIR AVAILABILITY.........■.................... 264 APPENDIX S: r e s u l t s o f t e s t s f o r s ig n if ic a n t d if f e r e n c e s in t h e n u m b e r o f 5 t r e e spe c ie s j BY SIZE CLASS FOUND AMONG 7 FOREST TYPES AT MOOSE FEEDING SITES ....................................................................L. . . 1 ................. 267 APPENDIX T: post hoc t e s t s d e t e r m in in g s ig n if ic a n t d if f e r e n c e s in t h e a v e r a g e p e r c e n t c o v e r AND HEIGHT OF SHRUB SPECIES AT MOOSE FEEDING SITES AMONG DIFFERENT AGES AND TYPES OF FORESTS ... 273 APPENDIX U : post hoc t e s t s d e t e r m in in g s ig n if ic a n t d if f e r e n c e s in f o r e s t st a n d CHARACTERISTICS AND AVERAGE NUMBER OF UNGULATE PELLET GROUPS AMONG 4 SUCCESSIONAL STAGES OF LODGEPOLE FORESTS .........................................................................................................................................................................278 APPENDIX V: r e s u l t s o f t e s t s f o r s ig n if ic a n t d if f e r e n c e s in p e l l e t g r o u p c o u n t s a n d TREE AND SHRUB CHARACTERISTICS AMONG 6 DISTANCES FROM THE BOUNDARY BETWEEN CLEARCUTS AND MATURE FORESTS.............................................................................................................................................................................286 APPENDIX W: r e s u l t s o f t e s t s f o r s ig n if ic a n t d if f e r e n c e s in a v er a g e n u m b e r o f b r o w s e d TWIGS, AVERAGE ANNUAL HEIGHT AND AVERAGE ANNUAL TWIGS PRODUCED AMONG 7 WILLOW COMMUNITIES, 7 WILLOW AND I BIRCH SPECIES, AND 7 YEARS (BETWEEN 1988 - 1997)...........................297 X LIST OF TABLES Table Page 1. Moose Horseback Index 1947-1949,1985-1992, and 1995-2001; dates of searches, number of moose observed, and age and sex of moose seen.............................................................................................................................. 24 2. Probability of at least one moose sighting per trip along the road between Gardiner and Cooke City, Montana, in each 2-month period computed empirically and from a logistic regression model.................................................. 27 3. Study unit location and sex of 14 radiocollared moose............................................ ............... 44 4. Percent of conifer cover types in 4 study units pre- and post-198 8 Yellowstone fires......................................................................................................................45 5. Cover categories (Mattson and Despain 1985) where moose were back-tracked to record browsed twigs..................................................................................... 55 6. Percent of moose diet represented by plant species comprising >5 percent of browsed stems during any period recorded before (1986-1988) and after (1988-1990) the 1988 fires and the early (November-December), mid- (January-February), and late (March-April) winter periods................................................................................................... 60 7. Percent of moose diet represented by plant species comprising >5 percent of browsed stems during any period recorded before (1986-1988) and after (1988-1990) the 1988 fires in the SB, YP, SC, and BC study units........................................................................................................... 61 8. Percent of twigs browsed by moose on burned and unbumed plants reported for all burned study units (excludes BC), each study unit, and the all (1988-1990), early (November-December), mid- (January-Februaiy), and late (March-April) winter periods post-fire............................61 9. Cover categories with > 4% of browsed stems in any period over four winters (1986-1990) in the early (November-December), mid- (January-February), and late (March-April) winter periods............................................ 62 10. Cover categories with > 4% of twigs browsed by moose during any period as expressed by a percentage of the total for the early (November-December), mid- (January-Februaiy), and late (March-April) winter periods before and after the 1988 fires (1986-1990).................................................. 63 11. Cover categories with > 4% of twigs browsed by moose during any period as expressed by a percentage of the total for the SB, YP, SC, and BC study units before (1986-1988) and after (1988-1990) the 1988 fires................................................................................................................................. 64 12. Five plant species with the most recorded twigs browsed by moose during the winter (1986-1990) in each cover category expressed as a percent of the total twigs browsed in that category.............................................................. 65 13. Distribution of browsed twigs among the early, mid-, and late successional stages of 3 types of coniferous forests...............................................................66 14. Average number of twigs browsed by moose per meter traveled in different cover categories during the first 2 winters (1988-89,1989-90) post-fire and the early (November-December), mid- (January -February), and late (March-April) periods of those winters.....................................................................66 15. Frequency of browsing on subalpine fir by size class.............................................................67 16. Average number of twigs browsed per subalpine fir by radiocollared and uncollared moose back-tracked through the snow. Data are reported as the number of browsed twigs/browsed trees for the pre- (1986-1988) and post­ fire (1988-1990) periods in all study units combined and each study unit............................. 68 17. Average number of twigs browsed per subalpine fir by radiocollared and uncollared moose back-tracked through the snow. Data are reported as the number of browsed twigs/browsed trees for the pre- (1986-1988) and post-fire (1988-1990) periods in each study unit.................................69 18. Cover type categories (Mattson and Despain 1985) used to characterize moose sightings along the Cooke City road........................................................................... 83 19. Cover categories (Mattson and Despain 1985) along fixed transects where moose tracks were searched fo r................................................................................... 84 20. Winter cover type use by 14 radiocollared moose summarized in 20 categories (Table 5). Data are reported for each study unit and represent all locations over 4 winters, November through April 1987-1991 ......................................... 88 21. Percent of radio collared moose (use) found in 13 cover categories (Table 20) compared to percent of cover categories available. Data are reported for the entire study area and represent all radio location points over 4 winters; November through April 1987-1999 pre- and post-1988 Yellowstone fires..................................................................................... 89 22. Winter cover type use by 14 radiocollared moose summarized in 20 categories (Table 5). Data are reported for each study unit and represent all radio location points over 4 winters; November through April 1987-1991 pre- and post-1988 Yellowstone fires..........................................................91 23. Percent of moose radio locations (use) found in 12 cover categories (Table 5) compared to percent of cover categories available. Data were recorded during the winters of 1986-87 and 1987-88 (pre-1988 Yellowstone fires) and are reported by study unit.................................................. 91 24. Percent of moose radio locations (use) found in 13 cover categories (Table 5) compared to percent of cover categories available. Data were recorded during the winters of 1988-89 and 1990-91 (post-1988 Yellowstone fires) and reported by study unit......................................................92 25. Comparison of pre- and post-fire radio collared moose use of 13 cover categories (Table 5) in 4 study units........................................................................................ 92 26. Percent of moose radio locations (use) found in burned and unbumed areas compared to percent of available fire affected areas.................... ................................ 93 27. Percentages of pellet groups found while back-tracking radiocollared and uncollared moose during the winter. Data are reported by cover category (Table 5) and the time frames of the early (November-December), mid- (January-Febmary), and late (March-April) periods of four winters (1986-1990) and the two years before (1986-1988) and after (1988-1990) the 1988 fires...................................................... 94 xi 28. Percentages of beds found while back-tracldng radiocollared and uncollared moose through the snow. Data are reported by cover category (Table 5) and the time frames of the early (November-December), mid- (January-February), and late (March-April) periods of four winters (1986-1990) and the two years before (1986-1988) and after (1988-1990) the 1988 fires.............................................95 29. Observations of moose along the Gardiner to Cooke City road, Yellowstone Park, summarized in 16 cover categories (Table 18)......................................... 96 30. Percent of moose tracks (use) found in 10 cover categories (Table 19) compared to percent of cover categories available. Data were recorded during the winters of 1986-87 and 1987-88 (pre-1988 Yellowstone fires)...........................100 31. Percent of moose tracks (use) found in 10 cover categories (Table 19) compared to percent of cover categories available. Data were recorded during the winters of 1988-89 and 1989-90 (post-1988 Yellowstone fires).........................101 32. Comparison of pre- and post-fire use by moose of 10 cover categories (Table 19) as indicated by tracks..........................................................................................101 33. Percent of elk tracks (use) found in 10 cover categories (Table 19) compared to percent of cover categories available. Data were recorded during the winters of 1986-87 and 1987-88 (pre-1988 Yellowstone fires). Early winter is November-December, mid-winter is January-?ebruary, and late winter is March-May........................................... 102 34. Percent of elk tracks (use) found in 10 cover categories (Table 19) compared to percent of cover categories available. Data were recorded during the winters of 1988-89 and 1989-90 (post-1988 Yellowstone fires)...................................................... ....................................... 103 35. Comparisons of pre- and post-fire use by elk of 10 cover categories (Table 19) as indicated by tracks.........................................................................................103 36. Selection by mule deer of 7 cover categories (Table 20) along BC transects as indicated by tracks............................................................................................. 104 37. Percent of moose tracks (use) found at various snow depths in the BC, SC, and SB study units compared to percent of the transect with various snow depths available............................................................................................... 118 38. Percent of elk tracks (use) found at various snow depths in the BC, SC, and SB study units compared to percent of the transect with various snow depths available............................................................................................... 120 39. Percent of mule deer tracks (use) found at various snow depths compared to percent of the transect with various snow depths available................................................................................................ 121 40. Cover type descriptions for moose feeding sites discovered while back-tracking moose.............................................................................................................. 129 41. Characteristics of 4 successional stages of lodgepole forests................................................. 129 42. Average percent canopy cover of deciduous shrub species in 7 forest types where moose browsing was recorded (Table 40): young (DF0), mid-age (DF2), and the oldest Douglas fir (DF3), young (LPl), mid-age (LP2), and the oldest lodgepole (LP3) and the oldest spruce-fir (SF)........................................................................................................................ 135 xii 43. Average height (cm) of 12 deciduous shrub species in 7 forest types where moose browsing was recorded (Table 40): young (DFO), mid-age (DF2), and the oldest Douglas fir (DF3), young (LPl), mid-age (LP2), and the oldest lodgepole (LP3), and the oldest spruce-fir (SF)........................................................................................................................ 136 44. Average age, DBH, number, and height of subalpine fir trees >5 m tall per 0.25 ha plot as well as number and m2 of subalpine fir patches in lodgepole forests of different ages (Table 40): LPO -0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance............................. ;............. ........ 138 45. Eight willow stands in SB and SC study units were used for sampling. Stands were distinguished as tall (>1.5 m) or low (<1.5 m) (Mattson and Despain 1985)................................................................................................. 168 46. Monthly snow conditions and presence of ungulate tracks, November-April, in 8 moose winter range willow stands, 1987-1991.................................172 47. Number of willow sampling plots at each site where plants were destroyed by the 1988 fires or drought/browsing stress compared with the number that reestablished by 1997 .........................................................................181 48. Number of plots of each willow species where plants were destroyed by the 1988 fires or drought/browsing stress compared with the number that reestablished by 1997 ............. :........................................................................ 182 49. Percent willow canopy cover and average height (m) in and out of the Slough Creek Exclosure, Absoraka-Beartooth Wilderness, 1963-1997................................ 183 50. Results of Newman-Keuls post hoc test determining significant differences among the 4 highest week averages of moose seen per-hour of observation at a SB willow stand over a 64-week period from April 1996 to June 1997.................................................... :......................................... 225 51. Number of moose observed from aircraft on the northern portion of the study area (north of the Yellowstone River)...................................................................227 52. Number of moose observed from aircraft on the southern portion of the study area (south of the Yellowstone River)...................................................................227 53. Results of Newman-Keuls post hoc test determining significant differences among the average moose home range size in 4 study units....................................................................................................................................... 229 54. Plant species appearing in the diet of moose over 4 winters, 1986-1991 .............................231 55. Plant species eaten by moose as determined by the percentage of the total number of browsed stems observed over 4 winters and for the early (November-December), mid- (January-Febraary), and late (March-April) winter periods over 4 years (1986-1990)......................................................233 56. Plant species in the diet of moose as determined by the percentage of the total number of browsed stems recorded before and after (1988-1990) the 1988 fires and the early (November-December), mid- (January-February), and late (March-April) winter periods...... .................................. 235 xiii 57. Plant species in the winter diet of moose as determined by the percentage of the total number of browsed stems recorded before (1986-1988) and after (1988-1990) the 1988 fires in the SB, YP, SC and BC study units................................................................................................................237 58. Percent of 5 tree species browsed and available as determined in 3-m radius plots at moose feeding sites in the early, mid-, and late successional stages of lodgepole pine forests.......................................................................239 59. Chi-square values from 5 tree species browsed and available as determined in 3-m radius plots at moose feeding sites in early, mid-, and late successional stages of lodgepole pine forests...................... .'........... i..................... 239 60. Bonferroni Z-statistic intervals for 5 tree species browsed and available as determined in 3-m radius plots at moose feeding sites in early, mid-, and late successional stages of lodgepole pine forests.......................................................... 239 61. Percent of 5 tree species browsed and available as determined in 3-m radius plots at moose feeding sites in early, mid-, and late successional stages of Douglas fir forests.................................................................................................. 240 62. Chi-square values from 5 tree species browsed and available as determined in 3-m radius plots at moose feeding sites in early, mid-, and late successional stages of Douglas fir forests...............................................................240 63. Bonferroni Z-statistic intervals for 5 tree species browsed and available as determined in 3-m radius plots at moose feeding sites in early, mid-, and late successional stages of Douglas fir forests............................................240 64. Percent of 5 tree species browsed and available as determined in 3-m radius plots at moose feeding sites in the late successional stage of spruce-fir forests...................................................................................................................241 65. Chi-square values and Bonferroni Z intervals from 5 tree species browsed and available as determined in 3-m radius plots at moose feeding sites in the late successional stage of spruce-fir forests...........................................241 66. Percent of browsed stems found in 20 cover categories over four winters (1986-1990) and the early (November-December), mid- (January-February), and late (March-April) winter periods ranked in descending order...................................................................................................243 67. Cover categories where stems browsed by moose were found expressed as a percentage of the total for the early (November-December), mid- (January-February), and late (March-April) winter periods before (1986-1988) and after (1988 -1990) the 1988 fires...................................................................................................245 68. Cover categories where stems browsed by moose were found during winter expressed as a percentage of the total for the SB, YP, SC, and BC study units before (1986-1988) and after (1988-1990) the 1988 fires........................................................................................................................................247 69. Results of Newman-Keuls post hoc test to determine significant differences in the average number of twigs browsed per subalpine fir in early, mid-, and late winter periods..............................................................................249 70. Results of Newman-Keuls post hoc test to determine significant differences in the average number of twigs browsed per subalpine fir pre-fire among study units................................................................................................ 249 xiv \ \ ■ 71. Results of Newman-Keuls post hoc test to determine significant differences in the average number of twigs browsed per subalpine fir post-fire among study units.............................................................................................. 249 72. Nutrient (proximate and elemental) analysis of subalpine fir summarized . for 4 winter periods, 1998-99, BC study un it..................... i................................................ 251 73. Nutrient (proximate and elemental) analysis of willow summarized for 4 winter periods, 1998-99, BC study unit....................................................................... 251 74. Nutrient (proximate and elemental) analysis of lodgepole pine summarized for 4 winter periods, 1998-99, BC study unit...................................................251 75. Nutrient (proximate and elemental) analysis of gooseberry summarized for 4 winter periods, 1998-99, BC study un it.......................................................................252 76. Nutrient (proximate and elemental) analysis of buffalobeny summarized for 4 winter periods, 1998-99, BC study un it.......................................................................252 77. Chi-square values from pre- and post-fire moose winter radiolocations in the study area and 20 cover categories............................................................................. 254 78. Bonferonni Z-statistic intervals for pre- and post-fire chi-square values from moose winter radiolocations in the study area and 20 cover categories......................254 79. Chi-square values from pre-fire moose winter radiolocations in 4 study units and 20 cover categories................................................................................................ 256 80. Bonferonni Z-statistic intervals for pre-fire chi-square values from moose winter radiolocations in 4 study units and 20 cover categories................................. 256 81. Chi-square values from post-fire moose winter radiolocations in 4 study units and 20 cover categories...................................................................................... 257 82. Bonferonni Z-statistic intervals for post-fire chi-square values from moose winter radiolocations in 4 study units and 20 cover categories.................................257 83. Chi-square values from pre- and post-fire moose winter radiolocations in 3 study units and areas with disparate fire effects............................................................259 84. Bonferonni Z-statistic intervals for post-fire chi-square values from moose winter radiolocations in 3 study units and areas with disparate fire effects...............................................................................................................259 85. Chi-square values from pre-fire winter encounters with moose tracks along fixed transects in 10 cover categories.................. ...................................................... 261 86. Bonferroni Z-statistic intervals for pre-fire chi-square values from winter encounters with moose tracks along fixed transects in 10 - cover categories.............................................................................. 261 87. Chi-square values from post-fire winter encounters with moose tracks along fixed transects in 10 cover categories.........................................................................261 88. Bonferonni Z-statistic intervals for post-fire chi-square values from winter encounters with moose tracks along fixed transects in 10 cover categories........................................ 262 89. Chi-square values from pre-fire winter encounters with elk tracks along fixed transects in 10 cover categories.......................................................................... 262 90. Bonferonni Z-statistic intervals for pre-fire chi-square values from winter encounters with elk tracks along fixed transects in 10 cover categories...................262 XV 91. Chi-square values from post-fire winter encounters with elk tracks along fixed transects in 10 cover categories.........................................................................263 92. Bonferonni Z-statistic intervals for post-fire chi-square values from winter encounters with elk tracks along fixed transects in 10 cover categories...................263 93. Chi-square values and Bonferroni Z-statistic intervals from winter encounters with mule deer tracks along fixed transects in 8 cover categories in the BS study un it.............................................................................................263 94. Chi-square values for percentages of moose tracks (use) found at various snow depths in the BC, SC, and SB study units compared to percent of the transect with various snow depths available..................................................265 95. Bonferroni Z-statistics intervals for moose tracks (use) found at various snow depths in the BC, SC, and SB study units compared to percent of the transect with various snow depths available.................................................. 265 96. Chi-square values for percentages of elk tracks (use) found at various snow depths in the BC, SC, and SB study units compared to percent of the transect with various snow depths available...............................................................265 97. Bonferroni Z-statistics intervals for elk tracks (use) found at various snow depths in the BC, SC, and SB study units compared to percent of the transect with various snow depths available...............................................................266 98. Chi-square values for percentages of mule deer tracks (use) found at various snow depths in the BC, SC, and SB study units compared to percent of the transect with various snow depths available..................................................266 99. Bonferroiii Z-statistic intervals for mule deer tracks (use) found at various snow depths in the BC, SC, and SB study units compared to percent of the transect with various snow depths available..................................................266 100. Average number of subalpine fir per 0.02-ha plot by size class at moose feeding sites in 7 forest types: young (DFO), mid-age (DF2), and oldest Douglas fir (DF3), young (LPl), mid-age (LP2), and oldest lodgepole (LP3) and oldest spruce-fir (SF) forests....................................................268 101. Average number of subalpine fir per 0.02-ha plot by size class at moose feeding sites in Douglas fir forests of different ages: young (DFO), mid-age (DF2), and oldest Douglas fir (DF3) forests...............................................268 102. Average number of subalpine fir per 0.02-ha plot by size class at moose feeding sites in lodgepole forests of different ages: young (LPl), mid-age (LP2), and oldest lodgepole (LP3) forests...................................................269 103. Average number of subalpine fir per 0.02-ha plot at moose feeding sites in late successional Douglas fir (DF3), lodgepole pine (LP3), and spruce-fir (SF) forests......... :..................... .................................................... .............. 269 104. Average number of subalpine fir per 0.02-ha plot by size class at moose feeding sites in early successional Douglas fir (DFO) and lodgepole pine (LPl) forests................................................................................................. 270 105. Average number of lodgepole pine per 0.02-ha plot by size class at moose feeding sites in 7 forest types: young (DFO), mid-age (DF2), and the oldest Douglas fir (DF3), young (LPl), mid-age (LP2), and the oldest lodgepole (LP3) and the oldest spruce-fir (SF) forests........................................270 xvi 106. Average number of spruce per 0.02-ha plot by size class at moose feeding sites in 7 forest types young (DFO), mid-age (DF2), and the oldest Douglas fir (DF3), young (LPl), mid-age (LP2), and the oldest lodgepole (LP3) and the oldest spruce-fir (SF) forests.........................................................271 107. Average number of Douglas fir per 0.02-ha plot by size class at moose feeding sites in 7 forest types young (DFO), mid-age (DF2), and the oldest Douglas fir (DF3), young (LPl), mid-age (LP2), and the oldest lodgepole (LP3) and the oldest spruce-fir (SF) forests.........................................................271 108. Average number of whitebark pine per 0.02-ha plot by size class at moose feeding sites in 7 forest types: young (DFO), mid-age (DF2), and the oldest Douglas fir (DF3), young (LPl), mid-age (LP2), and the oldest lodgepole (LP3) and the oldest spruce-fir (SF) forests........................................ 272 109. Average number of trees per plot 0.02-ha of the 5 tree species found in the study area for each of the 7 cover types where moose browsing occurred: young (DFO), mid-age (DF2), and the oldest Douglas fir (DF3), young (LPl), mid-age (LP2), and the oldest lodgepole (LP3) and the oldest spruce-fir (SF) forests....................................................................................272 110. Results of Newman-Keuls post hoc tests for significant differences in average percent canopy cover of ninebark among moose feeding sites in 5 forest types................................................................................................274 111. Results of Newman-Keulspost hoc tests for significant differences in average percent canopy cover of gooseberry among moose feeding sites in 7 forest types......... ;......................................................................... 274 112. Results of Newman-Keuls post hoc tests for significant differences in average height (cm) of gooseberry among moose feeding sites in 7 forest types.................................................................... ........................................ 274 113. Results of Newman-Keuls post hoc tests for significant differences in average percent canopy cover of grouse whortleberry among moose feeding sites in 4 forest types........................................................................274 114. Results of Newman-Keuls post hoc tests for significant differences in average height (cm) of grouse whortleberry among moose feeding sites in 4 forest types.....................................................................................275 115. Results of Newman-Keuls post hoc tests for significant differences in average percent canopy cover of buffaloberry among moose feeding sites in 7 forest types....................................................................................275 116. Results of Newman-Keuls post hoc tests for significant differences in average height (cm) of buffaloberry among moose feeding sites in 7 forest types .:i..........................................................................................................275 117. Results of Newman-Keulspost hoc tests for significant differences in average percent canopy cover of snowberry among moose feeding sites in 6 forest types....................................................................................275 118. Results of Newman-Keuls post Aoc tests for significant differences in average percent canopy cover of Rocky Mountain maple among moose feeding sites in 3 forest types........................................................................276 119. Results of Newman-Keulspost hoc tests for significant differences in average percent canopy coyer of serviceberry among moose feeding sites in 4 forest types......... .......................................................................... 276 xvii 120. Results of Newman-Keuls post hoc tests for significant differences in average percent canopy cover of honeysuckle among moose feeding sites in 5 forest types.................................................................................... 276 121. Results of Newman-Keuls post hoc tests for significant differences in average height (cm) of honeysuckle among moose feeding sites in 5 forest types................................................................................................. 276 122. Results of Newman-Keuls post hoc tests for significant differences in average height (cm) of huckleberry among moose feeding sites in 4 forest types.................................................................................................277 123. Results of Newman-Keuls post hoc test for significant differences in average age of 10 largest trees in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance.....................................................................................................................279 124. Results of Newman-Keuls post hoc test for significant differences in average number of moose pellet groups in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance.................................................................................................................... 279 125. Results of Newman-Keuls post hoc test for significant differences in average number of elk pellet groups in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance........................................ i.......................................................................... 279 126. Results of Newman-Keuls post hoc test for significant differences in average number of mule deer pellet groups in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2- 100 to 300, and LP3- 300+ years post-disturbance.................................................................................................................... 279 127. Average number of browsed and unbrowsed subalpine fir per 0.02-ha plot by size class in lodgepole forests of different ages: LPO -0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance............................280 128. Results of Newman-Keuls post hoc test for significant differences in average number of browsed subalpine fir <1 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years-post-disturbance...........................................................................................................280 129. Results of Newman-Keuls post hoc test for significant differences in average number of unbrowsed subalpine fir <1 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2- 100 to 300, and LP3- 300+ years post-disturbance................................................................................................................... 280 130. Results of Newman-Keuls post hoc test for significant differences in average percent browsed subalpine fir <1 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2- 100 to 300, and LP3- 300+ years post-disturbance.................................................................................... 281 xviii 131. Results of Newman-Keuls post hoc test for significant differences in average number of browsed and unbrowsed subalpine fir <1 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance................................................. . 132. Results of Newman-Keuls post hoc test for significant differences in average number of browsed subalpine fir 1-5 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance.................................................................................... 133. Results of Newman-Keulspost hoc test for significant differences in average number of unbrowsed subalpine fir 1-5 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance.................................................................................... 134. Results of Newman-Keuls post hoc test for significant differences in average percent browsed subalpine fir 1-5 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance.................................................................................... 135. Results of Newman-Keulspost hoc test for significant differences in average number of browsed and unbrowsed subalpine fir 1-5 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance.................................................. 136. Results of Newman-Keulspost hoc test for significant differences in average number of browsed subalpine fir <5 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2- 100 to 300, and LP3- 300+ years post-disturbance...... ,............................................................................ 137. Results of Newman-Keulspost hoc test for significant differences in average number of unbrowsed subalpine fir <5 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, andLP3- 300+ years post-disturbance.................................................................................... 138. Results of Newman-Keulspost hoc test for significant differences in average percent browsed subalpine fir <5 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance.................................................................................... 139. Results of Newman-Keuls post hoc test for significant differences in average number of browsed and unbrowsed subalpine fir <5 meter tall in 0.02-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2- 100 to 300, and LP3- 300+ years post-disturbance................................................. xix 281 281 2 8 1 2 8 2 2 8 2 2 8 2 282 2 8 3 2 8 3 XX. 140. Results of Newman-Keuls post hoc test for significant differences in average number of subalpine fir >5 meter tall in 0.02-ha plots among 4 successional stages of Iodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2- 100 to 300, and LP3- 300+ years post-disturbance..................................................... ........................ ..................................... 283 141. Results of Newman-Keuls post hoc test for significant differences in average number of subalpine fir in 0.25-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance..................................................................................................................... 283 142. Results of Newman-Keuls post hoc test for significant differences in average age of subalpine fir trees >5m in 0.25-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance..................................................................................................................... 284 143 . Results of Newman-Keuls post hoc test for significant differences in average DBH of subalpine fir trees >5m in 0.25-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance..................................................................................................................... 284 144. Results of Newman-Keuls post hoc test for significant differences in average number of subalpine fir trees >5m in 0.25-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance....................................................................................................................284 145. Results of Newman-Keuls post hoc test for significant differences in average height of subalpine fir trees >5m in 0.25-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2- 100 to 300* and LP3- 300+ years post-disturbance...........................284 146. Results of Newman-Keuls post hoc test for significant differences in average number of patches of subalpine fir trees in 0.25-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2- 100 to 300, and LP3- 300+ years post-disturbance................................................................................... ................................ 285 147. Results of Newman-Keuls post hoc test for significant differences in average size of patches (m2) of subalpine fir trees in 0.25-ha plots among 4 successional stages of lodgepole forests: LPO- 0 to 40, LPl- 40 to 100, LP2-100 to 300, and LP3- 300+ years post-disturbance.....................................................................................................................285 148. Average number of moose, elk, and deer pellet groups per 0.02-ha plot at different distances from a clearcut/forest boundary................................................... 287 149. Average percent canopy cover, per 0.02-ha plot of shrub species browsed by moose in clearcut and forest locations at different distances from the clearcut/forest boundary.......................................................................... 287 150. Results of tests for significant differences in percent canopy cover per 0.02-ha plot of shrub species browsed by moose among clearcut and forested locations at different distances from the clearcut/forest boundary................................................................................................................................287 151. Average percent canopy cover per 0.02-ha plot of shrub species browsed by moose in clearcuts and forests............................................................................288 152. Average height per 0.02-ha plot of shrub species browsed by moose in clearcut and forest locations at different distances from the clearcut/forest boundaiy.........................................................................................................288 153. Results of tests for significant differences in average height per 0.02-ha plot of shrub species browsed by moose among clearcut and forested locations at different distances from the clearcut/forest boundaiy................................................................................................................................288 154. Differences in average shrub height per 0.02-ha plot of shrub species browsed by moose between forests and clearcuts..................................................................289 155. Average number of browsed and unbrowsed subalpine fir per 0.02-ha plot in clearcut and forested locations at different distances from the clearcut/forest boundary.........................................................................................................289 156. Results of tests for significant differences in average number of browsed and unbrowsed subalpine fir per 0.02-ha plot in clearcut and forested locations at different distances from the clearcut/forest boundary..................................... 290 157. Differences in average number of browsed and unbrowsed subalpine fir per 0.02-ha plot in clearcut and forested locations...........................................................290 158. Average number of browsed and unbrowsed lodgepole per 0.02-ha plot in clearcut and forested locations at different distances from the clearcut/forest boundary.........................................................................................................291 159. Results of tests for significant differences in average number of browsed and unbrowsed lodgepole per 0.02-ha plot in clearcut and forested locations at different distances from the clearcut/forest boundary................................................................................................................................291 160. Differences in average number of browsed and unbrowsed lodgepole per 0.02-ha plot in clearcut and forested locations................................................................292 161. Average number of browsed and unbrowsed spruce per 0.02-ha plot in clearcut and forested locations at different distances from the clearcut/forest boundary.........................................................................................................292 162. Results of tests for significant differences in average number of browsed and unbrowsed spruce per 0.02-ha plot in clearcut and forested locations at different distances from the clearcut/forest boundary................................................................................................................................293 163. Differences in average number of browsed and unbrowsed spruce per 0.02-ha plot in clearcut and forested locations................................................................293 164. Average number of browsed and unbrowsed Douglas fir per 0.02-ha plot in clear cut and forested locations at different distances from the clearcut/forest boundary.........................................................................................................294 165. Results of tests for significant differences in average number of browsed and unbrowsed Douglas fir per 0.02-ha plot in clearcut and forested locations at different distances from the clearcut/forest boundary................................................................................................................................ 294 xxi 166. Differences in average number of browsed and unbrowsed Douglas fir per 0.02-ha plot in clearcut and forested locations.......................................................... 295 167. Average number of browsed and unbrowsed whitebark pine per 0.02-ha plot in clearcut and forested locations at different distances from the clearcut/forest boundary............................................ ............................................ 295 168. Results of tests for significant differences in average number of browsed and unbrowsed whitebark per 0.02-ha plot in clearcut and forested locations at different distances from the clearcut/forest boundary................................................................................................................................ 296 169. Differences in average number of browsed and unbrowsed whitebark pine per 0.02-ha plot in clearcut and forested locations.......................................................296 170. Average annual height per plot of 7 willow, I birch, and all species combined in centimeters........................................................................................................298 171. Results of Newman-Keuls post hoc test for significant differences in average annual height of Eastwood’s willow per 1-m plot among the years 1988-1991 and 1995-1997...................................................................298 172. Results of Newman-Keuls post hoc test for significant differences in average annual height of Farr’s willow per 1-m plot among the years 1988-1991 and 1995-1997.............................. i......................................... 298 173. Results of Newman-Keuls post hoc test for significant differences in average height of willow per 1-m plot among 7 willow communities......................................................................................................... i........ ...... 299 174. Average annual height of willow per 1-m plot in centimeters, all species combined, at each willow stand................................................................................299 175. Results of Newman-Keulspost hoc test for significant differences in average height of willow per 1-m plot among the years 1988-1991 and 1995-1997 at the Fisher Creek willow stand................... <............. . 299 176. Average annual utilization per 1-m plot for each species. Differences among means were determined using ANOVA or Kruskal-Wallis ANOVA by ranks..........................................................................................300 177. Results of Newman-Keuls post hoc test for significant differences in average annual percent of browsed willow twigs per 1-m plot of 7 willow and I birch species among the years 1988-1991 and 1995-1997 ....................................................................................................................... 300 178. Results of Newman-Keuls post hoc test for significant differences in average annual percent of browsed willow twigs per 1-m plot of Farr’s willow among the years 1988-1991 and 1995-1997........................................ 300 179. Results of Newman-Keulspost hoc test for significant differences in average annual percent of browsed willow twigs per 1-m plot of Drummond’s willow among the years 1988-1991 and 1995-1997..............................................................................................................................301 180. Results of Newman-Keuls post hoc test for significant differences , in average annual percent of browsed willow twigs per 1-m plot of Barclay’s willow among the years 1988-1991 and 1995-1997.............................................................................................................................. 301 xxii 181. Results of Newman-Keuls post hoc test for significant differences in average annual percent of browsed willow twigs per 1-m plot among 8 species.....................................................................................................301 182. Results of Newman-Keuls post hoc test for significant differences in average annual percent of browsed willow twigs per 1-m plot among 7 species.....................................................................................................302 183. Results of Newman-Keuls post hoc test for significant differences in average annual percent of browsed willow twigs per 1-m plot among 7 willow communities................................................................................302 184. Average annual utilization at each location...........................................................................302 185. Average number of twigs per plot without sites where willow did not persist............................................................................................................................... 303 186. Results of Newman-Keuls post hoc test for significant differences in average annual number of willow twigs per 1-m plot of 7 willow and I birch species among the years 1988-1991 and 1995-1997..............................................................................................................................303 187. Results of Newman-Keulspost hoc test for significant differences in average annual number of willow twigs per 1-m plot of Wolf’s willow among the years 1988-1991 and 1995-1997.............................................303 188. Results of Newman-Keuls post hoc test for significant differences in average annual number of willow twigs per 1-m plot of Booth’s willow among the years 1988-1991 and 1995-1997................................................ 304 189. Results of Newman-Keuls post hoc test for significant differences in average annual average number of willow twigs per 1-m plot of Farr’s willow among the years 1988-1991 and 1995-1997............................... 304 190. Results of Newman-Keuls post hoc test for significant differences in average annual number of willow twigs per 1-m plot of Drummond’s willow among the years 1988-1991 and 1995-1997 ..................................304 191. Results of Newman-Keuls post hoc test for significant differences in average annual number of willow twigs per 1-m plot of Eastwood’s willow among the years 1988-1991 and 1995-1997.........................................305 192. Average number of twigs per plot, including plots where willow did not persist...............................................................................................................................305 193. Results of Newman-Keuls post hoc test for significant differences in average annual average number of twigs per 1-m plot among the years 1988-1991 and 1995-1997..........................................................................305 194. Results of Newman-Keuls post hoc test for significant differences in average annual average number of willow twigs per 1-m plot of Wolf’s willow among the years 1988-1991 and 1995-1997..............................................................................................................................306 195. Results of Newman-Keuls post hoc test for significant differences in average annual number of willow twigs per 1-m plot of Farr’s willow among the years 1988-1991 and 1995-1997.............................................. 306 196. Results of Newman-Keuls post hoc test for significant differences in average annual number of willow twigs per 1-m plot of Drummond’s willow among the years 1988-1991 and 1995-1997................................... 306 xxiii xxiv 197. Results of Newman-Keuls post hoc test for significant differences in average annual e number of willow twigs per 1-m plot of Barclay’s willow among the years 1988-1991 and 1995-1997........................................ 307 198. Results of Newman-Keuls post hoc test for significant differences in average annual number of willow twigs per 1-m plot of Geyer’s willow among the years 1988-1991 and 1995-1997........................................... 307 199. Results of Newman-Keuls post hoc test for significant differences in average annual number of willow twigs per 1-m plot of Eastwood’s willow among the years 1988-1991 and 1995-1997......................................... 307 200. Results of Newman-Keuls post hoc test for significant differences in average annual number of twigs per 1-m plot of dwarf birch among the years 1988-1991 and 1995-1997................................................................308 XXV LIST OF HGURES Figure Page 1. Study area, study units, and the Northern Yellowstone Winter Range boundary..................................................................................................................................11 2. MooseHorsebackIndex 1947-1949,1985-1992, and 1995-2001; moose seen per day of searching............................................................................................ 25 3. Probability of seeing at least I moose while traveling the road from Gardiner, MT to Cooke City, MT (89 km) for each of the years 1987-1992 and 1995-1997...................................................................................................... 25 4. Probability of seeing at least one moose while traveling the road from Gardiner, MT to Cooke City, MT (89 km) by 2 month period for the years 1987-1992 and 1995-1997............ ........................................................:...................... 26 5. Probability of seeing at least I moose while traveling the 5 sections of road between Gardiner, MT, and Cooke City, MT, (177 km) before and after the 1988 fires for the years 1987-1992 and 1995-1997........................................... 26 6. Observations of moose during flights over SB and SC willow stands 1987-1990............................................................................................................................... 28 7. Ground observations of moose made daily every 1/2 hour during daylight hours at a SB willow stand, April 1996-June 1997. Average number seen per month and for each of the 64 weeks...........................................................29 8. Ground observations of moose made daily every 1/2 hour during daylight hours at a SB willow stand, April 1996-June 1997. Highest average number of moose seen per 1/2 hour each month......................................................30 9. Number of moose seen during flights of the NYWR and only north and south of the Yellowstone River from December 1988-May 1992 ........................ ..........30 10. Percent of 4 study units affected by the 1988 Yellowstone fires...........................................46 11. Percent decrease in conifer forests in 4 study units due to the 1988 Yellowstone fires..................................................... .............................................................. 46 12. The fate of 14 radiocollared moose in 4 study units and the number of months each survived after the 1988 Yellowstone fires.................................................... 47 13. Plant species that comprised >5% during any winter period of the total number of stems browsed by moose observed for the early (November-December), mid- (January-Febmary), or late (March-April) winter periods over 4 years (1-986-1990).......................................................59 14. Dry weight quantity (ppm) of calcium in 5 moose browse species summarized for 4 winter periods, 1998-99; Bear Creek Study U nit.....................................69 XXVl 15. Dry weight quantity (ppm) of potassium in 5 moose browse species summarized for 4 winter periods, 1998-99; Bear Creek Study U nit.....................................70 16. Dry weight quantity (ppm) of magnesium in 5 moose browse species summarized for 4 winter periods, 1998-99; Bear Creek Study U nit..................................... 70 17. Dry weight quantity (ppm) of sodium in 5 moose browse species summarized for 4 winter periods, 1998-99; Bear Creek Study U nit........ ............................ 71 18. Dry weight quantity (ppm) of phosphorus in 5 moose browse species summarized for 4 winter periods, 1998-99; Bear Creek Study U nit..................................... 71 19. Dry weight quantity (%) of ash in 5 moose browse species summarized for 4 winter periods, 1998-99; Bear Creek Study Unit..........................................................72 20. Dry weight quantity (%) of crude fiber in 5 moose browse species summarized for 4 winter periods, 1998-99; Bear Creek Study U nit.....................................73 21. Dry weight quantity (%) of crude protein in 5 moose browse species summarized for 4 winter periods, 1998-99; Bear Creek Study U nit.....................................73 22. Dry weight quantity (%) of ether extract in 5 moose browse species summarized for 4 winter periods, 1998-99; Bear Creek Study U nit..................................... 74 23. Winter cover type use by 14 radiocollared moose summarized in 20 categories (Table 5). Data are reported for the early (November-December), mid- (January-Febmary), and late (March-April) winter periods and represent all radio locations over 4 winters; November through April 1987-1991.....................................................................87 24. Winter cover type use by 14 radiocollared moose summarized in 20 categories (Table 5). Data are reported for the entire study area and represent all radio location points over 4 winters; November through April 1987-1991 pre- and post-1988 Yellowstone fires.......................................................... 89 25. Observations of moose during November and December along the Gardiner to Cooke City road, Yellowstone Park, summarized in 16 cover categories (Table 18)..................................................................................................... 97 26. Observations of moose during January and Febmary along the Gardiner to Cooke City road, Yellowstone Park, summarized in 16 cover categories (Table 18)......................................... 98 27. Observations of moose during March and April along the Gardiner to Cooke City road, Yellowstone Park, summarized in 16 cover categories (Table 18)............................................................................................................... 98 28. Observations of moose during May and June along the Gardiner to Cooke City road, Yellowstone Park, summarized in 16 cover categories (Table 18)................................................................................................................................ 99 29. Average available snow depth (cm) in the oldest lodgepole pine and the oldest spruce-fir forests during early (November-December)5 mid- (January-February) and late (March-April) winter compared with snow depths along routes used by browsing moose....................................................117 30. Average height of 11 shrub species recorded from plants browsed by moose in comparison with the average early (November-December)5 mid- (January-February)5 and late (March-April) winter snow depth in areas browsed by moose..................................................................................................Hg 31. Maximum5 minimum, and average snow depth (cm) in habitats used by and available to moose as determined by track encounters............................................119 32. Maximum5 minimum and average snow depth (cm) in habitats used by and available to elk as determined by track encounters..................................................120 33. Maximum5 minimum, and average snow depths (cm) in habitats used by and available to mule deer as determined by track encounters.......................................122 34. Average number of subalpine fir per 0.02-ha plot by height size class at moose feeding sites in 7 forest types (Table 40): young (DFO)5 mid-age (DF2), and the oldest Douglas fir (DF3), young (LPl)5 mid-age (LP2), and the oldest lodgepole (LP3) and the oldest spruce-fir (SF) forests (Appendix S5 Table 100)..................................................................131 35. Average number of lodgepole pine per 0.02 ha plot by height size class at moose feeding sites in 7 forest types (Table 40): young (DFO)5 mid-age (DF2), and oldest Douglas fir (DF3), young (LPl)5 mid-age (LP2), and oldest lodgepole (LP3) and oldest spruce-fir (SF) forests (Appendix S5 Table 105)..........................................................................................132 36. Average number of spruce per 0.02-ha plot by height size class at moose feeding sites in 7 forest types (Table 40): young (DFO)5 mid-age (DF2), and oldest Douglas fir (DF3), young (LPl)5 mid-age (LP2), and oldest lodgepole (LP3) and oldest spruce-fir (SF) forests (Appendix S5 Table 106)............ ............................................................................. 132 37. Average number of Douglas fir per 0.02-ha plot by height size class at moose feeding sites in 7 forest types (Table 40): young (DFO)5 mid-age (DF2), and oldest Douglas fir (DF3), young (LPl)5 mid-age (LP2), and oldest lodgepole (LP3) and oldest spruce-fir (SF) forests (Appendix S5 Table 107)......................................................................................................133 38. Average number of whitebark pine per 0.02 ha plot by height size. class at moose feeding sites in 7 forest types (Table 40): young (DFO)5 mid-age (DF2), and oldest Douglas fir (DF3), young (LPl)5 mid-age (LP2), and oldest lodgepole (LP3) and oldest spruce-fir (SF) forests (Appendix S5 Table 108)............................................ ......................................................... 134 xxvii .xxviii 39. Average number of trees per plot 0.02 ha of the 5 tree species found in the study area for each of the 7 cover types where moose browsing occurred (Table 40): young (DFO), mid-age (DF2), and the oldest Douglas fir (DF3), young (LPl), mid-age (LP2), and the oldest lodgepole (LP3) and the oldest spruce-fir (SF) forests (Appendix S, Table 109)......................................................................................................134 40. Average number of pellet groups of three ungulate species per 0.02 ha plots in lodgepole pine forests of different ages (Table 40): LPO -0 to 40, LPl- 40 to 100, LP2- 100 to 300, and LP3- 300+ years p o s t- d i s tu r b a n c e ..........................................................................................................................................................137 41. Average number of browsed and unbrowsed subalpine fir per 0.02 ha plot by height size class in lodgepole forests of different ages (Table 40): LPO -0 to 40, LPl- 40 tolOO, LP2- 100 to 300, and LP3- 300+years post-disturbance................................... ................................................... 137 42. Arrangement of study plots for measuring tree density in relationship to the clearcut/forest boundary and line transects for determining shrub canopy cover and height at each plot......................................... ............................... 147 43. Average number of moose, elk, and deer pellet groups per 0.02-ha plot at different distances from a clearcut/forest boundary. Sample plots were pa