Contents .

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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)
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