Contents Part I: INTRODUCTION CHAPTER 1 APPROACH: THE FLAMMULATED. BOREAL. AND GREAT GRAY OWL ASSESSMENTS Gregory D . Hayward GOALS .................................................................................................................................................................................................2 SCOPE OF ASSESSMENT .................................................................................................................................................................2 STANDARD FOR KNOWLEDGE .............................................................................................................................................3 LIMITATIONS ................................................................................................................................................................................... 3 DEFINITIONS ..................................................................................................................................................................................... 4 REFERENCES..................................................................................................................................................................................... 4 CHAPTER 2 METHODS AND TERMINOLOGY USED WITH STUDIES OF HABITAT ASSOCIATIONS D . Archibald McCallurn AN APPROACH TO ASSESSING HABITAT REQUIREMENTS .............................................................................................. 5 HABITAT REQUIREMENTS .............................................................................................................................................................5 HABITAT PREFERENCES ................................................................................................................................................................6 HABITAT SELECTION AND OCCUPANCY ...........................................................................................................................7 REFERENCES................................................................................................... Part 11: FLAMMULATED OWLS CHAPTER 3 CURRENT MANAGEMENT SITUATION: FLAMMULATED OWLS Jon Verner REFERENCES .................................................................................................................................................................................... 11 CHAPTER 4 REVIEW OF TECHNICAL KNOWLEDGE: FLAMMULATED OWLS D . Archibald McCallurn INTRODUCTION ............................................................................................................................................................................. 14 SYSTEMATICS ................................................................................................................................................................................. 14 Phylogenetic Position ......................................................................................................................................................14 Species Status ..........................................................................................................................................................................15 Subspecies and Geographic Variation .............................................................................................................................16 Fossil History .......................................................................................................................................................................... 16 DISTRIBUTION AND ABUNDANCE ......................................................................................................................................... 16 Recognized Distribution .................................................................................................................................................. 16 Breeding Range ......................................................................................................................................................................17 Nonbreeding Range ............................................................................................................................................................... 17 Estimates of Local Abundance and Population Trends ................................................................................................... 18 MOVEMENTS ................................................................................................................................................................................... 18 Migration .................................................................................................................................................................................18 Timing and Routes of Migration .........................................................................................................................................19 20 HABITAT OCCUPANCY ........................................................................................................................................................... Composite Requirements ..................................................................................................................................................... 20 A Hierarchical Scheme for Summarizing Information on Habitat ................................................................................. 20 Geographic Range ................................................................................................................................................................. 2 0 Biogeography of Related Species..................................................................................................................................... 20 Distinctive Features of the Owl's Range .................................................................................................................... 21 Climate and Thermoregulation ....................................................................................................................................... 21 Elevational Range .............................................................................................................................................................. 22 Landscape (Vegetation Type) ............................................................................................................................................... 22 General Floristics ...............................................................................................................................................................22 Characteristics of Entire Home Range ....................................................................................................................... 22 Home Range ...........................................................................................................................................................................23 Foraging Areas ................................................................................................................................................................... 23 Roost Sites ..........................................................................................................................................................................25 Microhabitat ...........................................................................................................................................................................2 5 Roost Sites ...........................................................................................................................................................................25 Singing Sites ........................................................................................................................................................................25 Foraging Sites and Maneuvers ...................................................................................................................... .................. 26 Characteristics of Nest Trees and Nest Cavities ..........................................................................................................26 Scale Effects ............................................................................................................................................................................. 27 Seasonal Difference in Habitat ............................................................................................................................................. 27 FOOD HABITS .................................................................................................................................................................................. 27 Feeding ....................................................................................................................................................................................27 Food Capture and Consumption ................................................................................................................................... 27 Diet ........................................................................................................................................................................................... 28 Major Food Items ..........................................................................................................................................................28 Role of Diet in Species Behavior .......................................................................................................................................... 29 BREEDING BIOLOGY ..................................................................................................................................................................... 29 Phenology of Courtship and Breeding ...........................................................................................................................29 Pair Formation .....................................................................................................................................................................30 Clutch Initiation and Laying ............................................................................................................................................ 30 Home Range Size .........................................................................................................................................................30 Courtship Characteristics .....................................................................................................................................................30 Pair Formation ....................................................................................................................................................................30 Courtship. Feeding. and Copulation .............................................................................................................................. 31 Clutch ......................................................................................................................................................................................31 Parental Care ..........................................................................................................................................................................31 DEMOGRAPHY ...............................................................................................................................................................................32 Life History Characteristics .................................................................................................................................................. 32 Age at First Reproduction .................................................................................................................................................32 Annual Fertility and Reproductive Success ..................................................................................................................32 First-Year Survival .......................................................................................................................................................33 Adult Annual Survival ......................................................................................................................................................33 Lifespan ...............................................................................................................................................................................34 Nonbreeders .......................................................................................................................................................................34 Geometric Rate of Natural Increase ...........................................................................................................................34 Ecologcal Influences on Survival and Reproduction .....................................................................................................35 Causes of Death ........................................................................................................................................................... 35 Social Pattern for Spacing ................................................................................................................................................ 35 Local Density Estimates .................................................................................................................................................. 36 Population Studies ............................................................................................................................................................. 36 Spot-Mapping and Equivalent Studies ......................................................................................................................... 36 Call Surveys .................................................................................................................................................................. 36 Limiting Factors ..................................................................................................................................................................... 37 Patterns of Dispersal .............................................................................................................................................................. 37 Natal Dispersal ................................................................................................................................................................... 37 Breeding Dispersal ............................................................................................................................................................. 38 Metapopulation Structure .................................................................................................................................................... 38 Unsaturated Habitat Hypothesis ..................................................................................................................................... 38 Suboptimal Habitat Hypothesis ...................................................................................................................................... 39 COMMUNITY ECOLOGY .............................................................................................................................................................. 39 Habitat Change and Vulnerability to Predation ................................................................................................................39 Competitors .........................................................................................................................................................................39 Complex Interactions ............................................................................................................................................................40 40 RESPONSE TO FOREST CHANGE .......................................................................................................................................... Stand Scale Response ........................................................................................................................................................40 Loggmg ................................................................................................................................................................................ 40 Fire ....................................................................................................................................................................................... 40 Population Response .............................................................................................................................................................41 Effects of Fragmentation ................................................................................................................................................... 4 1 Response to Human or Mechanical Disturbance ..........................................................................................................41 REFERENCES .................................................................................................................................................................................... 41 Chapter 5 Dynamics of Ponderosa and Jeffrey Pine Forests Penelope Morgan FOREST DESCRIPTION .............................................................................................................................................................47 Forest Extent ...........................................................................................................................................................................47 Landscapes. Communities. and Stands ..............................................................................................................................47 HUMAN USE OF FOREST RESOURCES ..................................................................................................................................... 50 Wood Products .......................................................................................................................................................................50 Grazing. Wildlife. Water. and Recreation Resources ........................................................................................................ 51 PRESETTLEMENT FORESTS .........................................................................................................................................................51 Paleoecology ........................................................................................................................................................................... 52 Historical Fire Regimes .........................................................................................................................................................52 53 Old-Growth Forests ......................................................................................................................................................... SUCCESSION AND DISTURBANCE ...........................................................................................................................................55 Fire as an Ecosystem Process ................................................................................................................................................55 Fire Effects...........................................................................................................................................................................56 Timber Harvest ....................................................................................................................................................................... 57 Insects ......................................................................................................................................................................................58 Dwarf Mistletoe and Other Pathogens ............................................................................................................................59 Other Disturbance .................................................................................................................................................................. 59 SUCCESSIONAL PATTERNS ....................................................................................................................................................59 Contrasting Patterns of Succession Following Logging and Fire ...............................................................................62 FUTURE VEGETATION PATTERNS ............................................................................................................................................ 63 Ecologcal Restoration ......................................................................................................................................................63 Changng Environments .......................................................................................................................................................64 INFORMATION NEEDS .................................................................................................................................................................64 65 REFERENCES .................................................................................................................................................................................... Chapter 6 Conservation Status of Flammulated Owls in the United States D . Archibald McCallum INTRODUCTION .............................................................................................................................................................................74 Are the Distribution and Abundance of the Flammulated Owl Declining in All or a Major Part of Its Range? .....74 Distribution ......................................................................................................................................................................... 74 Abundance ..........................................................................................................................................................................74 Do Habitats Vary in Their Capacity to Support Flammulated Owl Populations or to Support Particular Functional Activities of the Owl? ......................................................................................................................................................... 75 What Are the Important Characteristics of the High Quality Habitats? Do Habitats Vary in Their Capacity to Support Principal Prey Species? .............................................................................................................................................. 76 If the Flammulated Owl or Its Prey Rely on Particular Habitats. Are These Habitats Declining or Being Stressed by Current Management? ........................................................................................................................................................ 76 Global Climate Change ..................................................................................................................................................... 76 U . S. Forests Under Current Climatic Conditions ...................................................................................................77 Winter Range of Populations Breeding in the United States .......................................................................................77 Do the Life History and Ecology of the Flammulated Owl Suggest That Populations Are Vulnerable to Habitat Change? ...........................................................................................................................................................................7 7 Is a Conservation Strategy Needed for This Species? ...................................................................................................... 78 REFERENCES .................................................................................................................................................................................... 79 Chapter 7 Information Needs: Flammulated Owls D. Archibald McCallum INTRODUCTION .............................................................................................................................................................................80 STRENGTH OF EXISTING KNOWLEDGE TO FORMULATE A CONSERVATION STRATEGY ......................................80 Distribution ............................................................................................................................................................................8 0 Response of Flammulated Owls to Stand-Level Habitat Changes ...............................................................................80 Effects of Broad-Scale Habitat Changes on Movement Patterns ..................................................................................81 Relationship Between Foraging Behavior and Prey Abundance ....................................................................................82 Demography ...........................................................................................................................................................................82 Monitoring Methods ............................................................................................................................................................. 83 Dynamics of Primary Plant Communities Used by the Flammulated Owl ................................................................ 83 Historic Patterns of Distribution and Composition of Forest Communities Used by the Flammulated Owl .........83 A RESEARCH PLAN .......................................................................................................................................................................83 Outline of Suggested Research Projects .............................................................................................................................. 84 REFERENCES .................................................................................................................................................................................... 85 Part 111: BOREAL OWLS Chapter 8 Current Management Situation: Boreal Owls Jon Verner REFERENCES ....................................................................................................................................................................................89 Chapter 9 Review of Technical Knowledge: Boreal Owls Gregory D. Hayward INTRODUCTION ............................................................................................................................................................................. 92 SYSTEMATICS ............................................................................................................................................................................... 9 2 DISTRIBUTION ........................................................................................................................................................................... 9 3 Species Range ......................................................................................................................................................................... 93 North America ........................................................................................................................................................................ 93 94 Species Status and Trend ...................................................................................................................................................... MOVEMENTS: ANNUAL, SEASONAL, AND DAILY .......................................................................................................... 95 Annual Movements and Site Tenacity of Adults .......................................................................................................... 95 Dispersal Frequency ................................................................................................................................................... 96 Dispersal Distances ............................................................................................................................................................ 96 Annual Movements and Site Tenacity of Juveniles ..........................................................................................................97 Seasonal Movements ............................................................................................................................................................. 97 Movements Within the Home Range .................................................................................................................................. 97 Daily Movements ................................................................................................................................................................... 98 Diurnal Period ............................................................................................................................................................... 98 Nocturnal Period ................................................................................................................................................................ 98 HABITAT USE ................................................................................................................................................................................... 99 Broad Habitat Use Patterns ...........................................................................................................................................99 Landscape Scale Habitat Use ............................................................................................................................................... 99 Home Range Scale Habitat Use ...................................................................................................................................100 Microhabitat .......................................................................................................................................................................... 100 Nest Sites ........................................................................................................................................................................... 100 Nest Tree and Cavity Characteristics ............................................................................................................................ 101 Roost Sites .................................................................................................................................. ...................................101 Foragng Sites ...................................................................................................................................................................102 FOOD HABITS ................................................................................................................................................................................ 103 Feeding .................................................................................................................................................................................. 103 Foragng Movements ..................................................................................................................................................103 Prey Capture .....................................................................................................................................................................103 Diet ........................................................................................................................................................................................103 Quantitative Analysis ......................................................................................................................................................103 Seasonal Variation ............................................................................................................................................................ 105 Yearly Variation ................................................................................................................................................................ 105 Energetics ............................................................................................................................................................................ 106 Temperature Regulation ................................................................................................................................................... 1.07 Food Caches .......................................................................................................................................................................... 107 ECOLOGY OF PRINCIPAL PREY ............................................................................................................................................... 107 Red-Backed Vole .................................................................................................................................................................. 107 Non-Forest Voles .................................................................................................................................................................. 108 Deer Mouse ........................................................................................................................................................................... 108 Northern Flying Squirrel .................................................................................................................................................... 108 BREEDING BIOLOGY .................................................................................................................................................................. 109 Phenology of Courtship and Breeding ............................................................................................................................. 109 Courtship .......................................................................................................................................................................... 109 Nest Occupancy ............................................................................................................................................................... 109 Egg Laying ........................................................................................................................................................................109 Fledgmg ............................................................................................................................................................................. 109 Mating System and Sex Ratio .......................................................................................................................................109 Nest Site ................................................................................................................................................................................. 110 Nest .................................................................................................................................................................................... 110 Maintenance or Re-Use of Nests .................................................................................................................................... 110 Nesting .................................................................................................................................................................................. 110 Egg Laying and Care of Young ................................................................................................................................. 110 Growth and Development ........................................................................................................................................110 DEMOGRAPHY ............................................................................................................................................................................. 110 Life History Characteristics ................................................................................................................................................ 110 Age of First Reproduction .............................................................................................................................................. 110 Clutch ................................................................................................................................................................................111 Fledging Success and Population Productivity ........................................................................................................... 111 Lifetime Reproductive Success ...................................................................................................................................... 112 Proportion of Population Breeding .......................................................................................................................... 112 Survivorship ..................................................................................................................................................................... 112 Movements as Related to Demography and Metapopulation Structure ..................................................................... 113 Local Densities ..................................................................................................................................................................... 113 Spacing and Population Regulation .................................................................................................................................. 113 Spacing .............................................................................................................................................................................. 113 Territoriality ...................................................................................................................................................................... 116 Population Regulation .................................................................................................................................................... 116 COMMUNITY ECOLOGY ............................................................................................................................................................ 117 Predation on Boreal Owls ................................................................................................................................................... 117 Relationship With Prey Populations ....................................................................................................................... 117 Competitors ...................................................................................................................................................................... 117 BOREAL OWL RESPONSE TO FOREST CHANGE .......................................................................................................... 118 Nesting Habitat .................................................................................................................................................................... 118 Roosting Habitat .................................................................................................................................................................. 118 Foraging Habitat .................................................................................................................................................................. 118 Broad-Scale Habitat Change ........................................................................................................................................119 BOREAL OWL RESPONSE TO HUMAN OR MECHANICAL DISTURBANCE ............................................................. 119 ASSESSMENT OF SCIENTIFIC BASIS FOR PARTICULAR MANAGEMENT TOOLS ..................................................... 119 Monitoring ............................................................................................................................................................................119 Viability Analysis .................................................................................................................................................................120 Effects Criteria Identification .............................................................................................................................................120 Stand and Watershed Scale Silviculture Prescriptions .................................................................................................121 REFERENCES ..................................................................................................................................................................................121 Chapter 10 Dynamics of Subalpine Forests Dennis H. Knight INTRODUCTION ...........................................................................................................................................................................128 DISTRIBUTION AND STAND CHARACTERISTICS ............................................................................................................. 128 Comparing Subalpine and Boreal Forests ........................................................................................................................ 129 SUBALPINE FORESTS SINCE THE PLEISTOCENE ...............................................................................................................130 SPATIAL DISTRIBUTION OF SUBALPINE FORESTS ............................................................................................................. 130 SUBALPINE FOREST DYNAMICS .......................................................................................................................................131 Causes of Disturbance ......................................................................................................................................................... 131 Fire .....................................................................................................................................................................................131 Insects ................................................................................................................................................................................131 Disease ............................................................................................................................................................................... 131 Timber Harvesting ........................................................................................................................................................... 132 Contrasting Effects of Different Disturbances on Succession........................................................................................132 REFERENCES ..................................................................................................................................................................................135 Chapter 11 Conservation Status of Boreal Owls in the United States Gregory D . Hayward INTRODUCTION ...........................................................................................................................................................................139 Is the Distribution and Abundance of the Boreal Owl Declining in All or Part of Its Range? ................................. 139 Distribution .......................................................................................................................................................................139 Abundance .................................................................................................................................................................. 140 Do Habitats Vary in Their Capacity to Support Boreal Owl Populations or to Support Particular Activities of the Owl? What Are the Important Characteristics of Those Habitats? ..........................................................................140 140 Landscape .................................................................................................................................................................... Microhabitat ......................................................................................................................................................................141 Do Habitats Vary in Their Capacity to Support Principal Prey Species? ...................................................................141 If the Boreal Owl or Its Prey Select Particular Habitats. Are These Habitats Declining or Being Stressed by Current Management? ............................................................................................................................................................... 141 Do the Life History and Ecology of the Boreal Owl Suggest That Populations Are Vulnerable to Habitat 142 Change? .............................................................................................................................................................................. Cavity Nesting ..................................................................................................................................................................142 Productivity ......................................................................................................................................................................142 Survival ............................................................................................................................................................................. 143 Home Range Size .........................................................................................................................................................143 Trophic Position ..........................................................................................................................................................143 Metapopulation Structure .........................................................................................................................................144 WHAT IS THE CURRENT AND PROJECTED CONSERVATION STATUS OF THE BOREAL OWL? ............................ 144 WHAT ARE THE MANAGEMENT CONSIDERATIONS? .....................................................................................................145 146 REFERENCES .................................................................................................................................................................................. Chapter 12 Information Needs: Boreal Owls Gregory D . Hayward INTRODUCTION ........................................................................................................................................................................... 148 STRENGTH OF EXISTING KNOWLEDGE TO FORMULATE A CONSERVATION STRATEGY ....................................148 Distribution ........................................................................................................................................................................... 148 Response of Boreal Owls to Stand-Level Habitat Changes ........................................................................................... 148 Effects of Landscape-Scale Changes on Home Range Use ..........................................................................................149 Effects of Regional-Scale Habitat Changes on Movement Patterns ............................................................................ 149 Effects of Stand-Level Habitat Changes on Foraging Behavior .................................................................................... 149 Demography ......................................................................................................................................................................... 149 149 Dynamics of Primary Plant Communities ................................................................................................................. History of Distribution and Composition of Forest Communities ............................................................................ 150 A RESEARCH DIRECTION ..........................................................................................................................................................150 Integrate a Variety of Research Tools ................................................................................................................................150 Modeling ........................................................................................................................................................................... 150 Experimental Studies Using Management Treatments .........................................................................................151 Observational Studies .....................................................................................................................................................152 Beyond Tools ........................................................................................................................................................................152 Examine Boreal Owl Ecology at a Range of Scales .................................................................................................. 152 Investigate Geographic Variation ..................................................................................................................................152 Investigate Many Questions on the Same Site ......................................................................................................... 152 Small-Scale Demography ................................................................................................................................................ 153 REFERENCES .................................................................................................................................................................................. 153 Part IV: GREAT GRAY OWLS Chapter 13 Current Management Situation: Great Gray Owls Jon Verner REFERENCES .................................................................................................................................................................................. 158 Chapter 14 Review of Technical Knowledge: Great Gray Owls James R . Duncan and Patricia H . Hayward INTRODUCTION ........................................................................................................................................................................... 159 159 DISTINGUISHING CHARACTERISTICS .................................................................................................................................. SYSTEMATICS ................................................................................................................................................................................ 159 DISTRIBUTION AND ABUNDANCE ........................................................................................................................................ 160 North American Breeding Range ......................................................................................................................................160 North American Winter Range ..........................................................................................................................................160 Eurasian Range ..................................................................................................................................................................... 160 Abundance ............................................................................................................................................................................160 Population Trends ................................................................................................................................................................ 160 vii MOVEMENTS ................................................................................................................................................................................. 161 Sex Differences .....................................................................................................................................................................162 HABITAT USE ................................................................................................................................................................................. 163 Nesting Habitat .................................................................................................................................................................... 163 Nest Structure ................................................................................................................................................................... 163 Broad Scale Nesting Habitat ...........................................................................................................................................163 Home Range Area ................................................................................................................................................................165 Microhabitat .......................................................................................................................................................................... 165 Nesting .............................................................................................................................................................................. 165 Roosting ....................................................................................................................................................................... 165 Foragmg ............................................................................................................................................................................ 165 166 Winter Use ..................................................................................................................................................................... FOOD HABITS ................................................................................................................................................................................ 166 INFLUENCE OF PREY SPECIES BIOLOGY ON OWL MOVEMENTS ...............................................................................167 168 BREEDING BIOLOGY ................................................................................................................................................................... Chronology of Courtship and Breeding ..........................................................................................................................168 Courtship Songs ................................................................................................................................................................... 168 Parental Care ........................................................................................................................................................................ 169 DEMOGRAPHY ............................................................................................................................................................................. 169 Life History Characteristics ................................................................................................................................................ 169 Nesting Success ............................................................................................................................................................... 169 Survival .............................................................................................................................................................................169 Ecologcal Influences on Survival and Reproduction .................................................................................................... 170 Social Pattern for Spacing ...................................................................................................................................................170 Limiting Factors ................................................................................................................................................................ 171 Patterns of Dispersal of Young ........................................................................................................................................... 171 Characteristics of Non-Breeding Segment of Population .............................................................................................. 171 Metapopulation Structure .................................................................................................................................................. 171 COMMUNITY ECOLOGY ............................................................................................................................................................ 171 Predators ............................................................................................................................................................................... 171 Competitors .......................................................................................................................................................................... 172 REFERENCES ..................................................................................................................................................................................172 Chapter 15 Dynamics of Forest Communities Used by Great Gray Owls James R . Habeck INTRODUCTION ........................................................................................................................................................................... 176 NORTHERN ROCKY MOUNTAINS .......................................................................................................................................... 176 Phytogeographic Provinces in the Northern Rocky Mountains ...............................................................................176 Southern Continental Province ......................................................................................................................................179 Northern Continental Province ..................................................................................................................................... 183 Intermountain Province ................................................................................................................................................ 185 Inland Maritime Province .........................................................................................................................................188 Historical Changes in Rocky Mountain Vegetation ........................................................................................................ 189 OREGON CASCADES AND CALIFORNIA SIERRA NEVADA MOUNTAINS .................................................................191 Introduction ................................................................................................................................................................... 191 Central-Western And Southcentral Oregon Cascades ....................................................................................................192 Central-Western Cascade Range .................................................................................................................................192 Southcentral Cascade Range ..........................................................................................................................................192 California Sierra Nevada Range ........................................................................................................................................ 193 Historical Changes in Cascade and Sierra Nevada Vegetation .....................................................................................195 Summary: Relationships Between Forest Dynamics and Management of Great Gray Owls ................................... 195 Acknowledgments ............................................................................................................................................................... 196 REFERENCES .................................................................................................................................................................................. 196 ... Vlll Chapter 16 Conservation Status of Great Gray Owls in the United States Gregory D. Hayward INTRODUCTION .................................................................................................................................................................... 202 Is the Distribution and Abundance of the Great Gray Owl Declining in All or Part of Its Range? .........................202 Distribution .......................................................................................................................................................................202 Abundance ....................................................................................................................................................................2 0 2 Do Habitats Vary in Their Capacity to Support Great Gray Owl Populations or to Support Particular Activities of the Owl? What Are the Important Characteristics of Those Habitats? .......................................................................202 Do Habitats Vary in Their Capacity to Support Principal Prey Species? .................................................................... 203 If the Great Gray Owl or Its Prey Relies on Specific Habitats. Are These Habitats Declining or Being Stressed by Current Management? ..................................................................................................................................................... 203 Do the Life History and Ecology of the Great Gray Owl Suggest That Populations Are Vulnerable to Habitat Change? ............................................................................................................................................................................ 204 Platform Nesting .............................................................................................................................................................. 204 Productivity and Survival .............................................................................................................................................. 204 204 Home Range Size ............................................................................................................................................................. 204 Trophic Position ............................................................................................................................................................... What Is the Current and Projected Conservation Status of the Great Gray Owl? ..................................................... 204 WHAT ARE THE MANAGEMENT CONSIDERATIONS? .............................................................................................. 205 REFERENCES .................................................................................................................................................................................. 206 Chapter 17 Information Needs: Great Gray Owls Gregory D . Hayward INTRODUCTION ........................................................................................................................................................................... 207 STRENGTH OF EXISTING KNOWLEDGE TO FORMULATE A CONSERVATION STRATEGY ....................................207 207 Distribution ........................................................................................................................................................................... Response of Great Gray Owls to ........................................................................................................................................207 Stand-Level Habitat Changes ........................................................................................................................................... 207 Effects of Landscape-Scale Changes on Home Range Use ........................................................................................... 208 Effects of Regonal-Scale Habitat Changes on Movement Patterns ............................................................................208 Effects of Stand-Level Habitat Changes on Foraging Behavior ................................................................................ 208 Demography ........................................................................................................................................................................ 2 0 8 Dynamics of Primary Plant Communities .......................................................................................................................208 History of Distribution and Composition of Forest Communities ..........................................................................209 209 A RESEARCH PLAN ............................................................................................................................................................... Establish Research Goals ...............................................................................................................................................209 Develop Predictive Models ............................................................................................................................................ 209 Examine Mechanisms ..................................................................................................................................................... 210 Use Ecosystem Management ........................................................................................................................................210 Specific Research Topics ................................................................................................................................................ 210 REFERENCES.................................................................................................................................................................................. 211 APPENDIX A: About The Authors ..........................................................................................................................................212 ACKNOWLEDGMENTS.......................................................................................................................................................... 214 MAP 1. Flammulated Owl Locations and Distribution of Associated Vegetative Ecosystems in the United States (Envelope) MAP 2. Boreal Owl Locations and Distribution of Associated Vegetative Ecosystems in the United States ..... (Envelope) MAP 3. Great Gray Owl Locations in the United States ........................................................................................ (Envelope)