The Ecology of Large Mammals in Central Yellowstone: Sixteen Years of Integrated Field Studies

Cover_Large.jpeg......Page 1
Series Editors......Page 2
Copyright page.pdf......Page 3
TOT1.pdf......Page 4
TOT2.pdf......Page 5
TOT3.pdf......Page 6
Contributors to Volume 3.pdf......Page 7
Acknowledgments.pdf......Page 10
Integrated Science in the Central Yellowstone Ecosystem......Page 13
References......Page 21
The Central Yellowstone Landscape: Terrain, Geology, Climate, Vegetation......Page 24
Overview and Geographic Setting......Page 25
Geology......Page 27
Climate......Page 30
Wet and Dry Meadows......Page 33
Geothermal Area Vegetation......Page 34
Fire and Forests......Page 35
Mapping of Vegetation Type......Page 36
Mapping of Forest Cover......Page 37
Summary......Page 39
References......Page 41
Mapping Mean Annual Precipitation Using Trivariate Kriging......Page 43
Introduction......Page 44
Available Climate Station Data......Page 45
Kriging......Page 46
Kriging Variants as Applied to Estimation of Mean Annual Precipitation......Page 49
Trivariate Zonal Ordinary Kriging......Page 50
Other Developments......Page 52
Summary......Page 53
References......Page 56
Appendix......Page 57
Effects of Yellowstone's Unique Geothermal Landscape on Snow Pack......Page 59
Introduction......Page 60
Spatial Distribution of Geothermal Intensity......Page 63
Statistical Distribution of GHF......Page 67
Effect of GHF on Snow Pack......Page 68
Discussion......Page 69
References......Page 71
Effects of Wind, Terrain, and Vegetation on Snow Pack......Page 73
Introduction......Page 74
Terrain......Page 76
Snow Pack......Page 77
Sampling Design for Forest Effects......Page 78
Sampling Design for Terrain Effects......Page 79
Anemometers......Page 80
Statistical Methods......Page 82
Results......Page 83
Discussion......Page 84
References......Page 89
Modeling Spatial Snow Pack Dynamics......Page 91
Introduction......Page 92
Model History and Enhancements......Page 94
Model Structure and Inputs......Page 95
An "Incremental Random Sampling" Data Set......Page 96
Model Parameterization......Page 99
Model Testing......Page 106
Summary......Page 110
References......Page 113
Appendix......Page 115
Vegetation Dynamics of Yellowstone's Grazing System......Page 119
Introduction......Page 120
Remotely Sensed Data......Page 122
Field Sampling......Page 123
Extracting Phenology and Productivity Information from NDVI Time Series......Page 124
Phenological Patterns and Spatial Variations in Vegetation Performance......Page 126
Smoothed, Gap-Filled NDVI......Page 127
Patterns of Phenology and Productivity......Page 129
Discussion......Page 133
Summary......Page 135
References......Page 136
Elk Winter Resource Selection in a Severe Snow Pack Environment......Page 140
Introduction......Page 141
Data Collection......Page 142
Statistical Methods......Page 144
Results......Page 146
Discussion......Page 152
Summary......Page 156
References......Page 157
Diet and Nutrition of Central Yellowstone Elk During Winter......Page 160
Introduction......Page 161
Forage Quality......Page 162
Metabolizable Energy Intake......Page 164
Results......Page 166
Discussion......Page 172
Summary......Page 174
References......Page 175
Living in Yellowstone's Caldera: A Geochemical Trophic Cascade in Elk......Page 180
Introduction......Page 181
Methods......Page 182
Elk Sampling......Page 183
Results and Discussion......Page 184
References......Page 190
The Madison Headwaters Elk Herd: Stability in an Inherently Variable Environment......Page 193
Introduction......Page 194
Study Population......Page 196
Demographic Variables......Page 197
Vital Rate Covariates......Page 198
Model Development and Evaluation......Page 199
Constructing Population Models......Page 200
Results......Page 202
Discussion......Page 208
Summary......Page 214
References......Page 215
Partial Migration in Central Yellowstone Bison......Page 219
Introduction......Page 220
Development of Migration Response Variables......Page 221
Development of Covariates......Page 222
1970-1971 Through 2005-2006 Migration Patterns: Model Development and Statistical Analyses......Page 224
Temporal Variation in Migratory Behavior (1996-1997 Through 2005-2006)......Page 226
Historic Migration Patterns (1970-1971 Through 2005-2006)......Page 229
Estimates of Nutritional Intake......Page 230
Discussion......Page 231
Summary......Page 233
References......Page 234
Appendix......Page 237
Emigration and Density Dependence in Yellowstone Bison......Page 238
Yellowstone's Two Bison Herds......Page 239
Data Collection......Page 241
Population Models-1902-1954......Page 243
Population Models-1970-2000......Page 245
Population Models-1902-1954......Page 246
Population Models-1970-2000......Page 247
Discussion......Page 248
References......Page 252
Demography of Central Yellowstone Bison: Effects of Climate, Density, and Disease......Page 255
Introduction......Page 256
Demographic Variables......Page 257
Bison and Climate Covariates......Page 259
Model Development and Evaluation......Page 260
Results......Page 262
Discussion......Page 270
Summary......Page 273
References......Page 274
Appendix 14A.1......Page 277
Methods......Page 278
Results......Page 279
Wolf Recolonization of the Madison Headwaters Area in Yellowstone......Page 280
Reintroduction History......Page 281
Wolf Collaring and Handling......Page 283
Ground Monitoring on the Ungulate Winter Range......Page 284
Population Demographic Parameters......Page 285
Wolf Pack Territory Estimation......Page 286
Population Status......Page 287
Reproduction and Denning......Page 288
Survival, Mortality and Dispersal......Page 289
Spatial and Temporal Territoriality......Page 290
Discussion......Page 295
Summary......Page 298
References......Page 299
Wolf Prey Selection in an Elk-Bison System: Choice or Circumstance?......Page 301
Introduction......Page 302
Detecting and Identifying Wolf Kills......Page 305
Factors Influencing Wolf Prey Selection......Page 307
Prey Switching......Page 311
Results......Page 312
Factors Influencing Wolf Prey Selection......Page 314
Wolf Prey Switching with Murdoch's Equation......Page 318
Discussion......Page 320
Summary......Page 326
References......Page 327
Model Suite 3: Prey + Wolf Competition Models......Page 331
Model Suite 4: Prey + Snow Pack + Wolf Competition Models......Page 332
Wolf Kill Rates: Predictably Variable?......Page 334
Introduction......Page 335
Wolf Tracking and Kill Detection......Page 337
Kill Rate Estimation......Page 338
Evaluating Kill Rate Variation......Page 339
Evaluating the Functional Response for Elk......Page 340
Results......Page 342
Variations in Elk Kill Rate......Page 344
Variations in Bison and Total Kill Rates......Page 345
Functional Response......Page 347
Discussion......Page 351
Summary......Page 356
References......Page 357
Appendix A. A priori Model Lists for Evaluating Kill Rate Variation......Page 360
Appendix B. Description of Multiple Species Functional Response Models......Page 363
Spatial Responses of Elk to Wolf Predation Risk: Using the Landscape to Balance Multiple Demands......Page 365
Introduction......Page 366
Elk Winter Home Ranges......Page 368
Long-Distance Movements......Page 370
Covariates of Predation Risk......Page 371
Statistical Analyses......Page 372
A Priori Model Suites and Predictions......Page 373
Results......Page 374
Mixed Effects Linear Regression Results......Page 376
Dispersal and Migratory Movements......Page 378
Discussion......Page 380
Summary......Page 385
References......Page 386
Appendix......Page 390
Elk Group Size and Wolf Predation: A Flexible Strategy when Faced with Variable Risk......Page 392
Introduction......Page 393
Non-Wolf Predictors of Grouping Behavior......Page 395
Covariates of Predation Risk......Page 396
Statistical Analyses......Page 397
A Priori Model Suites and Predictions......Page 398
Temporal Trends in Elk Group Size......Page 399
Model Selection Results......Page 400
Discussion......Page 403
Summary......Page 410
References......Page 411
Elk Foraging Behavior: Does Predation Risk Reduce Time for Food Acquisition?......Page 414
Introduction......Page 415
Methods......Page 417
Landscape, Temporal, and Social Predictors of Elk Foraging Behavior......Page 419
Covariates of Predation Risk......Page 420
Statistical Analyses......Page 421
A Priori Model Suites and Predictions......Page 423
Results......Page 424
Model Selection Results......Page 426
Discussion......Page 430
References......Page 436
Appendix......Page 440
Changes in Elk Resource Selection and Distribution with the Reestablishment of Wolf Predation Risk......Page 442
Introduction......Page 443
Methods......Page 445
Results......Page 447
Discussion......Page 456
References......Page 464
Elk Nutrition after Wolf Recolonization of Central Yellowstone......Page 468
Introduction......Page 469
Methods......Page 470
Results......Page 472
Discussion......Page 473
Summary......Page 476
References......Page 477
Appendix......Page 479
The Madison Headwaters Elk Herd: Transitioning from Bottom-Up Regulation to Top-Down Limitation......Page 480
Introduction......Page 481
Demographic Variables......Page 483
Vital Rate Covariates, Predictions, Model Development, and Evaluation......Page 485
Constructing Population Models......Page 486
Adult Female Survival......Page 488
Elk Pregnancy Rates......Page 490
Calf Survival and Yearling Recruitment......Page 494
Population Estimates, Growth Rates and Projections......Page 495
Discussion......Page 499
References......Page 504
Apparent Competition and Regulation in a Wolf-Ungulate System: Interactions of Life History Characteristics, Climate, and Landscape Attributes......Page 509
Introduction......Page 510
Conceptual Wolf-Ungulate Models......Page 512
Apparent Competition......Page 515
Physical, Behavioral, and Environmental Vulnerability......Page 517
Predicting the Future of the Madison Headwaters Predator-Prey System......Page 521
Summary......Page 527
References......Page 528
Contrasting Wolf-Ungulate Interactions in the Greater Yellowstone Ecosystem......Page 531
Introduction......Page 532
Elk Populations and Study Site Descriptions......Page 534
Pregnancy Rate......Page 538
Elk Calf Recruitment......Page 539
Adult Female Survival......Page 540
Pregnancy Rate......Page 541
Elk Calf Recruitment in Relation to Condition, Drought, and Forage......Page 543
Adult Female Elk Survival......Page 547
Hunter Harvest and Estimated Wolf-Kill of Adult Female Elk......Page 548
Elk Population Trend......Page 550
Discussion......Page 552
Summary......Page 555
References......Page 556
Description of Study Areas......Page 559
Predator Numbers......Page 561
Pregnancy, early calf survival, and condition......Page 563
Elk Population Size and Composition......Page 564
Hunter Harvest......Page 565
Wolf Survival/Mortality......Page 566
Wildlife Responses to Park Visitors in Winter......Page 568
Introduction......Page 569
Methods......Page 570
Model Formulation......Page 571
Abundance, Demography, and Displacement of Wildlife......Page 572
Results......Page 573
Abundance, Demography, and Displacement of Wildlife......Page 577
Discussion......Page 579
Summary......Page 582
References......Page 583
Appendix 26A.1 Summary of Data from 25 Surveys (265 interactions) During Winter 2002-2003 That Were Inadvertently Omitted from Analyses......Page 585
Appendix 26A.2 Estimates of Abundance and Recruitment for Bison, Elk, Trumpeter Swans, and Bald Eagles Plotted Against Numbers of Visitors on Snowmobiles and Coaches During.........Page 586
Bison Winter Road Travel: Facilitated by Road Grooming or a Manifestation of Natural Trends?......Page 589
Introduction......Page 590
Analysis of Temporal Travel Patterns......Page 592
Analysis of Spatial Travel Patterns......Page 594
Temporal Variation in Bison Travel Behaviors......Page 595
Spatial Variation in Bison Road Travel......Page 596
Activity Patterns......Page 598
Discussion......Page 600
Summary......Page 603
References......Page 604
Appendix......Page 606
Effects of Snow and Landscape Attributes on Bison Winter Travel Patterns and Habitat Use......Page 608
Introduction......Page 609
Methods......Page 610
Snow Pack, Topography, and Habitat Type Covariates......Page 611
A Priori Hypotheses......Page 612
Model Development and Statistical Analyses......Page 613
Results......Page 614
Modeling and Predicting Spatial Variation in Bison Travel and Non-Travel Activities......Page 618
Discussion......Page 619
Summary......Page 627
References......Page 628
Appendix......Page 630
Communicating Ecological Knowledge to Students and the Public......Page 633
Approach and Technological Framework......Page 634
Simple Web-based "Data Mapper" Tool......Page 635
"Bison Ecology and Management" Undergraduate Education Module......Page 637
Bison Migration Virtual Interpretive Trail......Page 641
Upper Geyser Basin Virtual Interpretive Trails......Page 644
Electronic Field Trips-Windows Into Wonderland......Page 646
Decision Support Tool for Managers......Page 647
Summary......Page 649
References......Page 650
Science in Yellowstone: Contributions, Limitations, and Recommendations......Page 652
Expectations......Page 653
Contributions......Page 655
Limitations......Page 659
Recommendations......Page 662
Summary......Page 664
References......Page 665
Index.pdf......Page 670