iS* IT , r » (. TRANSACTIONS OF THE LINNAEAN SOCIETY OF NEW YORK Volume VIII Development of Behavior in Precocial Birds By Margaret Morse Nice New York July, 1962 THE LINNAEAN SOCIETY OF NEW YORK The Linnaean Society of New York, organized in 1878, is the second oldest of existing American ornithological societies. Regular meetings of the Society are held on the second and fourth Tuesdays of each month from September to May, inclusive. Informal meetings are held at least once a month during June, July and August All meetings (except the annual dinner meeting on the second Tuesday of March) are open to the public and are held at the American Museum of Natural History. Persons interested in natural history are eligible for election to membership in the Society. Annual dues for Active Members are $6.00, and for Associate Members are $2.50. The Society conducts field trips and maintains a circulating library for its members. 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QW \ /O/-/ TRANSACTIONS / / OF THE LINNAEAN SOCIETY OF NEW YORK Volume VIII Development of Behavior in Precocial Birds By Margaret Morse Nice Illustrated with Line Drawings by the Author New York July, 1962 To Albert Hochbaum without whom this book would never have been written PREFACE For four seasons—from 1951 to 1954—my daughter Constance and I were privileged to study the early development of precocial chicks at the Delta Waterfowl Research Station at Delta, Manitoba. The primary object was to compare the development of behavior in precocial birds with that in altricial birds—a subject on which I had concentrated earlier. Those were rich experiences at Delta beside the great marsh and its wildlife, so vividly described by Albert Hochbaum (1944). The friendly, eager atmosphere of staff and students, engaged in the study of waterfowl life, and the visits of biologists from the Old and New Worlds were most stimulating. We are deeply indebted to Albert Hochbaum, Director of the Station, for his generous welcome to us and his unfailing assistance in our problems. We owe much to Peter Ward, Superintendent of the Hatchery, to Norman Godfrey, and to many of the fellow students who brought us eggs of different species and showed us nests they had found. Helen Hays was especially helpful. Fruitful consultations were held with Frank McKinney, Nicholas Collias, and especially William H. Elder, whose help was invaluable in working out a scheme of classification of birds with eight categories from the most precocial to the altricial, namely, Table 2. I sketched our subjects directly from life, sometimes in pencil, sometimes in ink. (As might be expected, the scale is not always con- sistent.) In this activity Albert Hochbaum gave me excellent criticisms and suggestions. Working up of the results of our studies was delayed by other projects: an excursion into development of mammalian behavior (Nice et al., 1956) and especially an investigation of the history of statements on incubation periods from Aristotle to the present day (Nice, 1954a, 1954b). Both these lines of research shed light on the studies at Delta. When I was able to turn my whole attention to these, I found that the original subject had broadened. Having discovered so many analogies between the behavior development of precocial and altricial birds and between them and mammals, it seemed wise to devote an introductory chapter to the functions of parental care throughout the Animal Kingdom. Here Konrad Lorenz, visiting this country in 1954, gave me excellent suggestions, as did Lars von Haartman two years v —; later. The scientist to whom I am most indebted for the validity of Chapters I and II is the late Karl P. Schmidt, Director of Zoology at the Chicago Natural History Museum, whose broad knowledge, espe- cially of herpetology, enabled him to criticize what I had collected from the literature and to give me additional information. Many ornithologists have helped through reading and criticizing portions of the manuscript : Chapters I through IV—Emmet R. Blake, John Emlen, Louise de Kiriline Lawrence, S. Charles Kendeigh, Austin L. Chapters I Rand ; through V and Chapter XIV—George A. Bar- tholomew, Jr.; Chapters V and XIV—Thomas R. Howell; Chapter VI, Section A Albert Hochbaum, Frank McKinney Chapter VI — ; Milton Weller; Chapter VII—Derek Goodwin, Eckhard Hess; Chap- ter VIII, Section A K. E. L. Simmons and most of the chapters — ; Laidlaw Williams. To all of these friends I am deeply grateful from ; each one I received help, even though I did not follow all of their suggestions. Special thanks are due to Anne Wachenfeld and Eugene Eisenmann for their editorial labors. To my daughter Janet I am indebted for the typing of the manu- script. To Constance my debt is great indeed she took the major part ; of the care of the baby birds ; she shared the watching, and often contributed illuminating insights into the meaning of various actions of our subjects. I am extremely grateful to the Wildlife Management Institute of Washington, D. C., sponsors of the Delta Waterfowl Research Station, for financial assistance in publishing this study. A word as to the organization of this volume. It is a study of behavior development in animals with chief emphasis on the Class Aves, particularly on representative precocial birds. The first four chapters are introductory. They include: a brief survey of the functions of parental care and of its occurrence throughout the Animal Kingdom a summary of some aspects of behavior development in five classes of classification according to maturity at hatching vertebrates ; a of birds and a discussion and comparison of stages in behavior development of an altricial bird, a mammal, and a precocial bird. Chapter V is con- cerned with megapodes, the most precocial of all birds. Chapters VI through XI are devoted to the birds studied by us, mostly at Delta, and Chapter XII to generalizations founded on these studies. Chapters XIII and XIV are largely based on research—anatomical and physio- logical—of other workers. VI CONTENTS Preface v List of Illustrations xii I Functions of Parental Care 1 A. Parental Care in Invertebrates 2 B. Parental Care in Vertebrates 3 1. Fishes. 2. Amphibians. 3. Reptiles. 4. Birds. 5. Mammals. C. Summary 12 II Examples of Post-Embryonic Development of Behavior in Vertebrates 13 A. Observations on Embryonic Behavior 13 B. Some Aspects of Development in Seven Vertebrates (Table 1) 13 C. Summary 17 III Precox and Altrix in Birds 18 A. Maturity at Hatching in Birds (Table 2) 18 B. Maturity at Hatching Throughout the Class Aves (Table 3) 19 C. Percentage of Yolk in Eggs of Precocials and Altricials (Table 4) 25 D. Summary 26 IV Stages of Development in Behavior 28 A. Five Stages of Development in Altricial Birds 28 B. Comparison of an Altricial Bird and Mammal (Table 5) 29 C. Comparison of an Altricial and a Precocial Bird (Table 6) 31 D. Summary 34 vii ; ; ; ; ; ; ; ;: V Precocial Chicks Independent of Their Parents: Mega- podes 35 A. The Eggs and Their Incubation 35 B. The Chicks 36 1. The Brush Turkey. 2. The Mallee Fowl. 3. Comparison with Other Precocial Birds. C. Summary 40 VI Precocial Chicks That Follow Parents and Find Their Own Food: Ducks, Killdeer, Spotted Sandpiper 41 A. Ducks 42 1. Numbers and Species Watched (Table 7). 2. Ex- amples of Post-hatching Behavior. 3. Development of Behavior in Ducklings. 4. Stage II—Preliminary (Table 8). 5. Stage III—Transition. 6. Stage IV —Locomotory : a. Eating. 7. Stage V— Socializa- tion : a. Aggression b. Bathing c. Flight. 8. Escape Reactions. 9. Bond to Parent. 10. Bond to Brood Mates. B. Killdeer 55 1. Examples of Post-hatching Behavior. 2. Develop- ment of Behavior in Killdeer. 3. Stage II— Prelim- inary (Table 9). 4. Stage III—Transition. 5. Stage IV—Locomotory: a. Play-fleeing b. /ump -and- flap c. Scratching the head. 6. Stage V—Socialization a. Aggression b. Bathing c. Foot-patting d. Flying e. Other coordinations. 7. Escape Reactions. 8. Bond to Parents. C. Spotted Sandpiper 66 1. Example of Post-hatching Behavior. 2. Develop- ment of Behavior in Spotted Sandpipers (Table 10). 3. Stage II—Preliminary. 4. Stage III—Transition. 5. Stage IV—Locomotary : a. Stalking prey b. Play- fleeing. 6. Stage V—Socialization: a. Aggression; b. Flight. 7. Stage VI—Independence : a. Food- washing. 8. Escape Reactions. 9. Bond to Parent and Brood Mates. D. Summary 74 vm ; ; ; ; ; ; ; ; VII Precocial Chicks that Follow Parents and Are Shown Food: Domestic Chicken 76 A. Domestic Chickens 76 1. Example of Post-hatching Behavior. 2. Develop- ment of Behavior in Chickens (Table 11). 3. Stage II—Preliminary. 4. Stage III—Transition: a. Ex- ploratory pecking b. Scratching the head. 5. Stage IV—Locomotory : a. Comfort movements b. Scratch- ing the ground c. Sun-bathing d. Rushing away with food. 6. Stage V—Socialization : a. Play- fighting b. Dust-bathing c. Flying d. Play-fleeing e. Pecking each other. 7. Escape Reactions. 8. Re- sponses to Parents. 9. Bond to Brood Mates. B. Summary 91 VIII Precocial Chicks That Follow Parents and Are Fed by Them: Grebes, Virginia Rail, Sora, Coot 92 A. Grebes 92 1. Examples of Post-hatching Behavior.