U.S. Fish and Wildlife Service Migration of Birds Circular 16 Migration of Birds Circular 16 by Frederick C. Lincoln, 1935 revised by Steven R. Peterson, 1979 revised by John L. Zimmerman, 1998 Division of Biology, Kansas State University, Manhattan, KS Associate editor Peter A. Anatasi Illustrated by Bob Hines U.S. FISH & WILDLIFE SERVICE D E R P O A I R R E T T M N EN I T OF THE U.S. Department of the Interior U.S. Fish and Wildlife Service TABLE OF CONTENTS Page PREFACE..............................................................................................................1 INTRODUCTION ................................................................................................2 EARLY IDEAS ABOUT MIGRATION............................................................4 TECHNIQUES FOR STUDYING MIGRATION..........................................6 Direct Observation ....................................................................................6 Aural ............................................................................................................7 Preserved Specimens ................................................................................7 Marking ......................................................................................................7 Radio Tracking ..........................................................................................8 Radar Observation ....................................................................................9 EVOLUTION OF MIGRATION......................................................................10 STIMULUS FOR MIGRATION......................................................................13 WHEN BIRDS MIGRATE ..............................................................................16 Time of Year..............................................................................................16 Time of Day ..............................................................................................19 FLIGHT SPEED AND RATE OF MIGRATION........................................24 MIGRATORY FLIGHT ALTITUDE..............................................................30 SEGREGATION DURING MIGRATION ....................................................32 As Individuals or Groups of Species......................................................32 By Age ......................................................................................................34 By Sex........................................................................................................35 GEOGRAPHIC PATTERNS OF MIGRATION ..........................................37 Populations Within Species ....................................................................37 Short Distance Migration ......................................................................38 Long Distance Migration........................................................................40 iii ORIENTATION AND NAVIGATION ..........................................................42 Vector Navigation ....................................................................................42 Orientation Cues ......................................................................................43 INFLUENCE OF WEATHER........................................................................46 INFLUENCE OF TOPOGRAPHY ................................................................49 PERILS OF MIGRATION ..............................................................................51 Storms........................................................................................................51 Aerial Obstructions..................................................................................51 Exhaustion ................................................................................................52 ROUTES OF MIGRATION..............................................................................53 General Considerations ..........................................................................53 Flyways and Corridors............................................................................53 Narrow Routes ........................................................................................54 Converging Routes ..................................................................................57 Principal Routes From North America ................................................61 Atlantic Oceanic Route ............................................................................61 Atlantic Coast Route and Tributaries ..................................................61 Mackenzie Valley-Great Lakes-Mississippi Valley Routes and Tributaries ................................................................................65 Great Plains-Rocky Mountain Routes ..................................................67 Pacific Coast Route ..................................................................................68 Pacific Oceanic Route ..............................................................................72 Arctic Routes ............................................................................................72 PATTERNS OF MIGRATION ........................................................................73 Loops..........................................................................................................73 Dog-legs ....................................................................................................78 Pelagic Wandering....................................................................................81 Leap-frogging ..........................................................................................83 Vertical Migration ....................................................................................83 Pre-migratory Movements ....................................................................84 Vagrant Migration....................................................................................84 FUTURE DIRECTIONS ................................................................................87 BIBLIOGRAPHY ..............................................................................................90 LIST OF BIRD SPECIES MENTIONED IN TEXT ..............................109 iv PREFACE TO THE 1998 EDITION Frederick C. Lincoln’s “Migration of Birds” was published in 1935. Lincoln’s writing style effectively communicated the wonders of bird migration to a wide audience, both young and old, experienced observers of birds as well as the simply curious. Indeed the demand for this little book was so great that it was revised in 1950 and soon was out of print again. In 1979, Steven R. Peterson developed a second revision, adding additional examples and presenting an understanding of bird migration that reflected current research. The style, figures, and most of the content of the origi- nal publication were retained, but new illustrations were added where appropriate. In this present revision large sections of the text have remained unchanged from the previous revision or only slightly modified to make the discussion compatible with current understanding. The geographic emphasis of Lincoln and the wealth of pertinent examples added by Peterson have been maintained. I have made substantial changes, however, in sections dealing with the evolution of migration, stimulus for migration, orientation and navigation, and the influence of weather. I have also changed the emphasis of the final section to reflect current concerns. While some investigators are mentioned by name, specific studies are not cited in the text. An exten- sive bibliography has been included for those interested in pursuing the subject further. I have relied heavily upon the bibliographies on migration research prepared by Stanley H. Anderson and Loren W. Ayers, University of Wyoming Fish and Wildlife Coop Unit and Thomas S. Litwin, as well as the bibliography in Peterson’s revision. Additional citations have been suggested by Daniel R. Petit and Stephanie L. Jones, U.S. Fish and Wildlife Service. This edition was due to the support of the Fish and Wildlife Service Regional Nongame Migratory Bird Coordinators: Tara Zimmerman, Kent Wohl, Steve Lewis, Daniel Petit, Diane Pence, Stephanie Jones, Bill Howe, and Richard Coon. I am indebted to all these investiga- tors and most grateful for their assistance. John Zimmerman, 1998 1 INTRODUCTION The changing picture of bird populations throughout the year intrigues those who are observant and who wish to know the source and destination of these birds. While many species of fish, mammals, and even insects undertake amazing migratory journeys, birds as a group are the most mobile creatures on Earth. Even humans with their many vehicles of loco- motion do not equal some birds in mobility. No human population moves each year as far as from the Arctic to the Antarctic with subsequent return, yet Arctic Terns do. Birds are adapted in their body structure and physiology to life in the air. Their feathered wings and tails, bones, lungs and air sacs, and their meta- bolic abilities all contribute to this amazing faculty. These adaptations make it possible for birds to seek out environments most favorable to their needs at different times of the year. This results in the marvelous phenom- enon we know as migration––the regular, recurrent, seasonal movement of populations from one geographic location to another and back again. Throughout human experience, migratory birds have been