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Handbook of Bird Biology

Kenneth P. Able, Ph.D. Dr. Stephen W. Kress
John Alcock, Ph.D. Dr. Donald E. Kroodsma
George A. Clark, Ph.D. Kevin J. McGowan, Ph.D.
Marie Eckhardt Daniel Otis
Howard E. Evans, Ph.D. Sandra Podulka
Alan Feduccia N. John Schmitt
John W. Fitzpatrick, Ph.D. Stanley A. Temple, Ph.D.
J. B. Heiser, Ph.D. David W. Winkler, Ph.D.





Rick Bonney
Director of Program Development and Evaluation

Rick has a B.S. in wildlife biology and a M.P.S. in natural resources extension, both from Cornell University, and he has been working at the Lab since 1981. A Cornell Senior Extension Associate, his research focuses on development and evaluation of inquiry-based science education projects.


Sandy Podulka
Education Associate

The primary editor of the course, Sandy also developed the course exams. She received a B.S. in Wildlife Biology from Cornell University and an M.S. in Zoology (Animal Behavior) from the University of Maryland, where she studied the function of song repertoires in Song Sparrows.



Ron Rohrbaugh
Acting Director of the Conservation Science Program

Second of the course editors and currently Acting Director of the Conservation Science Program, Ron guided the production of the course. He holds B.S. and M.S. degrees in Wildlife Biology from Penn State University. Ron's research focuses on the breeding biology of American Kestrels, grassland ecology, and using citizen science to study and monitor bird populations.


Kenneth P. Able, SUNY Albany
CHAPTER 5: Birds on the Move: Flight and Migration

Kenneth P. Able is Professor of Biology in the Department of Biological Sciences at the State University of New York in Albany, where he has been a member of the faculty since 1971. He received his B.S. and M.S. degrees from the University of Louisville, and his Ph.D. from the University of Georgia. Ken's research focuses on bird migration, particularly the mechanisms of orientation and navigation.

Ken has been passionately interested in birds since childhood. He has birded extensively across North America and in Mexico, Costa Rica, Peru, Spain, France, Germany, Italy, Switzerland, the United Kingdom, Australia, New Zealand, and South Africa. His favorite destination is Australia, because the avifauna is unique and the species fantastic.


John Alcock, Arizona State University
CHAPTER 6: Understanding Bird Behavior

John Alcock is Regents' Professor of Biology at Arizona State University, Tempe. John is currently researching the evolution of insect mating systems, with a special emphasis on the diversity of male mating tactics among bee species native to the Sonoran Desert. Earlier research has focused on birds, however, and his textbook, Animal Behavior, An Evolutionary Approach uses many bird examples to illustrate all aspects of the modern study of behavior.

John began bird watching at age 5 (bird number one was the mallard), and seeing a good bird still boosts his heart rate. In attempts to keep his heart rate up, he has visited Costa Rica, Ecuador, Argentina, Australia, and several countries in Europe.


George A. Clark, University of Connecticut
CHAPTER 3: Form and Function: The External Bird

George A. Clark, Jr. is Professor Emeritus of Ecology and Evolutionary Biology at The University of Connecticut in Storrs. He received his Ph.D. in Biology from Yale University, where he specialized in ornithology. After spending two years at the University of Washington in Seattle, he moved to The University of Connecticut where he spent 32 years as a faculty member.

His 200 publications reflect his research interests in the structure, behavior, distribution, and evolution of birds. He is past president of the Association of Field Ornithologists, served as Co-Editor of the book Perspectives in Ornithology/ Essays Presented for the Centennial of the American Ornithologists' Union, and has led educational field trips for groups to observe birds in North and South America, Europe, and Africa.

A birder since his high school days in Pennsylvania, George now resides in Vermont and enjoys seeking birds by walking or snowshoeing in the northern New England hills.


Marie Z. Eckhardt, Cornell University
Birds and Humans: A Historical Perspective

Marie Eckhardt is a biologist, formerly employed with the Education Department of the Cornell Lab of Ornithology. Her academic background in vertebrate zoology and experience as a museum consultant and as collections manager at the New York State Museum provided a strong foundation for her varied contributions at the Lab. Marie has been interested in animals and the natural world for as long as she can remember—an interest she credits to early and regular museum visits.


Howard E. Evans, Cornell University
CHAPTER 4: What's Inside: Anatomy and Physiology

Howard E. Evans received both his B.S. and Ph.D. in comparative anatomy from Cornell University, where he became a faculty member in the Veterinary College in 1950. There he taught gross anatomy of the horse and cow for seven years, and anatomy of the dog, bird, and fish for 36 years. He was secretary of the college for 12 years and served as chairman of Anatomy from 1976 to 1986, when he retired. He continues to teach a course on the literature and materials of natural history, and to lecture in several other courses.

Howie's research concerned the anatomy of reptiles and birds, the replacement of teeth in fishes, the plant-induced cyclopis in sheep, fetal development of the dog, and anatomy of tropical fishes. His most recent works include the third edition of Miller's Anatomy of the Dog (1993), the fifth edition of Guide to the Dissection of the Dog (2000 with Dr. de Lahunta), and the third edition of Anatomy of the Budgerigar and Other Birds (1996). He is co-editor of the Handbook of Avian Anatomy published by the Nuttall Ornithological Club of Harvard University. He has served as president of both the American and The World Association of Veterinary Anatomists, and was an associate editor of the American Journal of Anatomy and the Journal of Morphology.

A native of New York City, Howie and his wife Erica have led natural history trips for the Cornell Adult University to the Virgin Islands; Hawaii; Sapelo Island, Georgia; East and South Africa; Papua, New Guinea; and Antarctica. He has lectured in China, Russia, Taiwan, Hungary, Switzerland, Germany, England, Thailand, Brazil, Mexico, and Japan.


Alan Feduccia, University of North Carolina
Evolution of Birds and Avian Flight

Alan Feduccia is S. K. Heninger Professor of Biology at the University of North Carolina at Chapel Hill, where he has been for 30 years. He received his Ph.D. at the University of Michigan, where his thesis focused on the evolution of woodhewers and ovenbirds.

Alan's career has focused on vertebrate evolution, the evolution of birds, and the tempo and mode of the evolution of modern groups of birds. His interest in the origin of birds blossomed in the late 1970s when he wrote The Age of Birds for Harvard University Press. In 1973 he wrote a rebuttal to the theory of hot-blooded dinosaurs in the Journal of Evolution, and since that time has been involved in the debate on bird origins. His latest book, The Origin and Evolution of Birds (Yale University Press, 1999) treats many of the main issues in the bird evolution controversy, and takes what he calls the "ornithological" position, that is, that birds evolved flight from the trees down, and from a common ancestor with dinosaurs, but not directly from them.

Alan's interest in birds began as a teenager. Later, as an undergraduate student at Louisiana State University, he had the good fortune to participate in "bird" expeditions to Honduras, El Salvador, and Peru. He has maintained an interest in Neotropical birds ever since.


John W. Fitzpatrick, Cornell University
CHAPTER 10: Bird Conservation

John W. Fitzpatrick (Ph.D., Princeton University, 1978) is Director of the Cornell Laboratory of Ornithology, where he arrived in September 1995. He was Executive Director of Archbold Biological Station, a private ecological research foundation in central Florida, from 1988 through Aug. 1995, and was Curator of Birds at the Field Museum of Natural History (Chicago) from 1978 to 1989. He is a Fellow of the American Ornithologists' Union (AOU). In 1985 the AOU awarded him its highest research honor (Brewster Award), for his book and numerous research articles (co-authored with Glen Woolfenden) on demography, social behavior, and conservation of the endangered Florida Scrub-Jay.

Fitzpatrick has led many expeditions to remote areas of South America, especially the western Amazonian basin and the Andean foothills. He has published numerous papers on Neotropical birds, including descriptions of seven bird species new to science. Co-Author of the book, Neotropical Birds: Ecology and Conservation (Univ. Chicago Press, 1996), he has been engaged in applying science to real world conservation issues throughout his career. Most recently, he helped design and implement a major network of ecological preserves in central Florida by convening panels of scientific experts and by engaging county, state, and federal agencies, nongovernmental organizations, and private industry in the process.

He serves on the national governing boards of The Nature Conservancy, the National Audubon Society, and the Center for Biodiversity and Conservation at the American Museum of Natural History. He is on two Endangered Species Recovery Teams, including that of the world's rarest bird, the Hawaiian Crow. He enjoys watercolor painting, has been a birdwatcher since kindergarten.


J. B. Heiser, Cornell University
CHAPTER 4: What's Inside: Anatomy and Physiology

J. B. Heiser is a Senior Lecturer in the Department of Ecology and Evolutionary Biology at Cornell University. A vertebrate evolutionary ecologist, J. B. received his B.S. from Purdue Univerisity and his Ph. D. from Cornell University. For the past 35 years has taught a variety of courses in vertebrate comparative anatomy and ecology.

J. B. began teaching on the Ithaca campus, but soon was teaching field courses at the Shoals Marine Laboratory, an isolated island facility in the Gulf of Maine that is cooperatively run by Cornell University and the University of New Hampshire. Eventually he became director of the Shoals program, a position that he held for 15 years. He won the Clark Award for distinguished teaching and has traveled to every continent and ocean to teach natural history "on location." Although J. B. was trained as a marine biologist and his research focuses on the evolution and interrelationships of coral reef fish, he has considerable natural history experience in tropical forests worldwide. In his travels he has watched birds (and fish) in every major biome and biogeographic region that the planet has to offer, but he still gets a thrill out of backyard birding in Upstate New York.


Stephen W. Kress, Audubon
CHAPTER 2: A Guide to Bird Watching

Stephen W. Kress is Vice President for Bird Conservation for the National Audubon Society, Manager of the Society's Maine Coast Seabird Sanctuaries, and Director of the Seabird Restoration Program's Project Puffin. He also is ornithology program director for the Audubon Camp in Maine, an adjunct faculty member in the Wildlife Department at the University of Maine, Orono, and a Visiting Fellow at the Cornell Lab of Ornithology, where each spring he teaches a popular course called Spring Field Ornithology.

As director of Audubon's Seabird Restoration Program, Steve advises and manages the development of techniques for re-establishing various Maine seabird colonies, such as Atlantic Puffins, Leach's Storm-Petrel, and Arctic, Common, and Roseate Terns. In the Pacific region, he has studied the role of vocalizations in attracting endangered Dark-rumped Petrels to artificial burrows in the Galápagos Islands, and Short-tailed Albatross to decoys on Midway Island. He is author of many books, including The National Audubon Society's Birder's Handbook, The Bird Garden, and Project Puffin, as well as the Golden Guide to Bird Life. He also has authored numerous scientific papers on seabird biology and conservation.

During most of the year Steve lives on 33 acres of woods and meadows near Ithaca, New York, where he manages his land for songbirds and works on methods for restoring populations of Northern Bobwhite Quail. He spends summers on the Maine coast, continuing his lifelong interest in restoring nesting seabird colonies.


Donald E. Kroodsma, University of Massachusetts
CHAPTER 7: Vocal Behavior

Birds enter our lives in different ways. Some of us have been bird-crazy as long as we can remember, but others discovered birds later. I was a late-comer, as birds grabbed me from the chemistry lab during my last year of college. How grateful I am that they have never let go. Immediately after college I took two summer bird courses from the famed Olin Sewall Pettingill, who was then director of the Cornell Lab of Ornithology but taught at the University of Michigan field station in Pellston, Michigan. He put a tape recorder, headphones, and parabolic microphone in my hands and told me to "go out and tape record some birds." In doing so, he changed my life. I went to graduate school at Oregon State to study how young wrens learn their songs; and then for eight years I had the good fortune to work with Peter Marler at Rockefeller University in New York, on all aspects of bird song. The time since college has been spent reveling in the who, what, when, where, how, and why of bird song.


Kevin J. McGowan, Cornell University
CHAPTER 1: Introduction: The World of Birds

Kevin J. McGowan is the instructor of the Home Study Course in Bird Biology and the new online course Courtship and Rivalry in Birds at the Cornell Laboratory of Ornithology. He received his B.S. and M.S. degrees in Zoology from the Ohio State University and a Ph.D. in Biology from the University of South Florida, where he studied the social development of Florida Scrub-Jays. He came to Cornell University in 1988 as curator of the Ornithology and Mammalogy collections in the Department of Ecology & Evolutionary Biology. In addition to caring for the collections he conducted research on crows and taught classes in specimen preparation, field collecting, the relationships of birds, and Neotropical canopy biology. He moved to the Lab in 2001.  He helped create the original All About Birds web site and wrote the Bird Guide section. He co-edited and wrote much of The Second Atlas of Breeding Birds in New York State.

Kevin's primary research focuses on behavioral ecology of birds. Currently he is studying the reproductive and social behavior of two crow species in central New York state and investigating the impact of West Nile virus on crow populations. He is an Elected Member of the American Ornithologists' Union, director and webmaster for the Federation of New York State Bird Clubs, and a member of the New York State Avian Records Committee. He was former Secretary of the Ornithological Societies of North America (OSNA), and former editor of the Ornithological Newsletter, a bi-monthly OSNA publication.

An avocational birder since childhood, Kevin has traveled throughout North America and to Europe, Africa, and Central and South America to watch and study birds.


Daniel Otis, Cornell University
Birds and Humans: A Historical Perspective

Daniel Otis grew up on a farm in upstate New York. Since 1988 he has worked as a freelance editor, proofreader, and writer for Living Bird and other Lab of Ornithology projects. Currently a Ph.D. student in horticulture at Cornell, his research focuses on conservation of the world's maple species and on assessing the extent and effects of Norway maple invasiveness on northeastern forests.

Although plants rather than birds became his main interest, his appreciation of nature derives partly from memorable experiences with birds: "Once when I was a teenager, on a stormy November day, as I walked along a hedgerow," he writes, "I was astonished to come across an isolated little tree crowded with silent birds I'd never seen before—Cedar Waxwings. Too exhausted to fly, they merely shuffled down their perches a bit when I came close. And in March of some years, the air was alive day and night with the sound of immense flocks of Canada geese flying between the lake and the muddy cornfields around our house." Experiences such as these, he notes, quicken one's appreciation of the vivacity and fascination of the natural world, and help us understand what conservationists are fighting for.


Sandra G. Podulka, Cornell University
Birds and Humans: A Historical Perspective

Sandy Podulka received a B.S. in Wildlife Biology from Cornell University and an M.S. in Zoology (Animal Behavior) from the University of Maryland, where she studied the function of song repertoires in Song Sparrows. After graduate school, Sandy was a Research Technician at Cornell for Dr. Stephen T. Emlen, analyzing the social behavior of White-fronted Bee-eaters, but discovered she wanted to spend more time sharing her love of nature with others, so turned toward environmental education. She worked at the Cayuga Nature Center for several years, and as an Adjunct Professor of Biology at Tompkins Cortland Community College from 1988 to 1996, teaching courses in biology and conservation. Since 1986, she has worked at the Cornell Lab of Ornithology, in research, writing, public education, and editingÛmost recently as one of the editors of the Home Study Course in Bird Biology.

Sandy spent her childhood knee-deep in muddy ponds trying to catch tadpoles and frogs, and roaming fields collecting butterflies—her first real love. She has enjoyed birds as long as she can remember, but they did not take center stage until she took a summer ethology course at Cornell University from Dr. Bill Dilger. He hauled his students out before dawn every morning and taught them to recognize birds by their songs, and Sandy has been listening to them ever since. She has participated in Christmas Bird Counts and Breeding Bird Atlases, and has traveled to Costa Rica, Belize, Trinidad, and Peru to watch birds, but her favorite bird watching site is her yard—which overlooks a beaver pond with an ever-changing cast of avian actors.


N. John Schmitt

N. John Schmitt is a wildlife illustrator with a lifelong interest in birds. Since leaving the United States Army in 1973 John has devoted his life to a variety of ornithology related endeavors which have taken him to many countries in Latin America, Asia, and Europe. He has worked as a field biologist for both the Peregrine Falcon and California Condor recovery programs.

John's work as a field biologist led to more serious devotion to illustrating birds. He has illustrated several books, including the National Geographic Society's 3rd edition of Birds of North America; Clark's A Field Guide to the Raptors of Europe, the Middle East, and North Africa; and Skutch's Birds Asleep. Currently, he is working on bird field guides for Peru and India. John is a self-taught taxidermist/museum preparator whose work is on display in several California museums. He also co-leads ecotours in the United States and abroad.

A native of California, John is an avid bird watcher. His time in the field provides valuable inspiration and is integral to maintaining his enthusiasm. He considers his notebooks, which he fills with written and sketch notes, as a source of his most valuable reference and inspiration. John encourages everyone interested in natural history to keep a notebook.


Stanley A. Temple, University of Wisconsin
CHAPTER 9: From Individuals to Ecosystems: Understanding Bird Ecology

Stanley A. Temple is the Beers-Bascom Professor of Conservation in the Department of Wildlife Ecology at the University of Wisconsin in Madison. He is also the Chair of the graduate program in Conservation Biology and Sustainable Development in the Institute for Environmental Studies at Madison. A quintessential Cornellian, Stan earned his B.S., M.S., and Ph.D. all at Cornell, and for years has served as a member of the Lab of Ornithology's Administrative Board.

Stan's professional activities focus on avian ecology and bird conservation, with a special emphasis on endangered species. He and his students have worked with some of the world's most endangered birds, including Peregrine Falcons, California Condors, Whooping Cranes, and dozens of endangered endemic species on islands around the world. To date, none of those species has become extinct, and most are doing significantly better as a result of his work.

Stan has been interested in birds as long as he can remember. Birds of prey have always been among his favorites, and he has been a falconer for 45 years. He feels fortunate to have incorporated all of his ornithological pleasures into his professional life.


David W. Winkler, Cornell University
CHAPTER 8: Nests, Eggs, and Young: The Breeding Biology of Birds

David Winkler was born and raised in Sacramento, California. Unlike the rest of his family, he was a naturalist from about age four. After a progression of enthusiastic interest in butterflies, wildflowers, and herps, he finally settled on birds in his early teens. He learned and studied local birds alone for two years, when, at his first Christmas Count, he ran into Rich Stallcup. David then spent the last two years of high school learning a great deal from Rich about the birds of California. While in high school, an American Birds article introduced David to The Herring Gull's World by Niko Tinbergen, and a senior English project on gull taxonomy followed within a couple of years. By his freshman year at U. C. Davis, David was dreaming about all the species he might some day study as opposed to seeing for his list. While attending U. C. Davis, David and friends secured National Science Foundation funding for an ecological study of Mono Lake. This cemented his attachment to the Mono Basin and its bird life, and his friendship with David Gaines, with whom he co-founded the Mono Lake Committee in 1978. A dissertation on the clutch sizes of California Gulls (with Frank Pitelka at Berkeley) at Mono and Great Salt Lakes was followed by post-docs at University of Gothenburg, Sweden (with Malte Andersson); Oxford University (with John Krebs); and Cornell University (with Paul Sherman). David joined the Cornell faculty in 1988 upon the retirement of Tom Cade, and his research on swallows world-wide, and the Ithaca Tree Swallows in particular, has thrived ever since.