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Ducks, Geese, and of the World by Paul A. Johnsgard Papers in the Biological Sciences

2010

Ducks, Geese, and Swans of the World: Contents, Preface, & Introduction

Paul A. Johnsgard University of Nebraska-Lincoln, [email protected]

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Johnsgard, Paul A., "Ducks, Geese, and Swans of the World: Contents, Preface, & Introduction" (2010). Ducks, Geese, and Swans of the World by Paul A. Johnsgard. 2. https://digitalcommons.unl.edu/biosciducksgeeseswans/2

This Article is brought to you for free and open access by the Papers in the Biological Sciences at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Ducks, Geese, and Swans of the World by Paul A. Johnsgard by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. DUCKS, GEESE, and SWANS of the World Paul A. Johnsgard Revised Edition Ducks, Geese, and Swans of the World

By Paul A. Johnsgard

The only one-volume comprehensive survey of the available in English, this book combines lavish illustration with the most recent information on the natural history, current distribution and status, and identification of all the .

After an introductory discussion of the ten tribes of Anatidae, separate accounts follow for each of the nearly 150 recognized species. These include scientific and vernacular names (in French, German, and Spanish as well as English), descrip- tions of the distribution of all recognized subspecies, selected weights and mea- surements, and identification criteria for both sexes and various age classes. The “Natural History” section of each species account considers habitats, foods, so- cial behavior, and reproductive biology. Finally, a short discussion of the species’ present population status and current evidence about its evolutionary relation- ships are presented. A glossary of technical terms and derivations of vernacular names and a reference list of more than three hundred sources supplement the text.

A large number of illustrations—more than 170 line drawings and 132 distribu- tion maps—were prepared especially for this volume, which also includes 59 color plates. Each species is illustrated in either line drawing or photograph or both.

One of the world’s foremost authorities on waterfowl behavior, Paul A. Johns- gard is emeritus professor of biological sciences at the University of Nebraska– Lincoln. He is the author of more than 40 books, including Handbook of Waterfowl Behavior, Waterfowl: Their Biology and Natural History, Cranes of the World, and Quails of , North American of Upland and Shoreline, Waterfowl of North America, Birds of the Great Plains, Birds of the Rocky Mountains, North American , and Quetzals of the World, Hawks, Eagles, and Falcons of North America, The of the World, The Avian Brood Parasites: Deception at the Nest, The of North America, Arena Birds: Sexual Selection and Be- havior, and dozens more.

Cover photograph of whooper swans by Paul A. Johnsgard

Ducks, Geese, and Swans of the World

First published and copyright © 1978 by the University of Nebraska Press.

Electronic edition published 2010 by the University of Nebraska–Lincoln Libraries. Copyright 2010 by Paul A. Johnsgard. All rights reserved

Library of Congress Cataloging in Publication Data

Johnsgard, Paul A. Ducks, geese, and swans of the world.

Bibliography: p. 387. Includes index. 1. Anatidae. I. Title. QL696.A52J62 598.4’ 1 Dedicated to The Wildfowl Trust Whose efforts have helped us to understand and preserve these magnificent birds

Contents

Page Page List of Illustrations xi Hawaiian 74 76 Preface xv Goose 80 Introduction to the Family Anatidae xvii 83 Tribe Anseranatini () Red-breasted Goose 86 Magpie Goose 3 Tribe Cereopsini () Tribe Dendrocygnini (Whistling or Tree Ducks) Cape Barren Goose 93 Spotted Whistling 7 Tribe Stictonettini (Freckled Duck) Plumed 8 Freckled Duck 99 10 Tribe Tadornini (Sheldgeese and ) 12 Blue-winged Goose 103 15 104 White-faced Whistling Duck 17 Magellan Goose 105 Cuban Whistling Duck 20 Goose 108 Black-bellied Whistling Duck 21 Ashy-headed 109 White-backed Duck 24 Ruddy-headed Sheldgoose 111 Tribe Anserini (Swans and True Geese) 113 Mute 29 115 31 Ruddy 116 Black-necked Swan 34 Cape Shelduck 118 36 120 39 Shelduck 122 Whistling Swan 42 124 Bewick Swan 44 Northern (Common) Shelduck 126 47 129 49 Tribe Tachyerini (Steamer Ducks) Bean Goose 52 Flying 135 White-fronted Goose 55 Magellanic Flightless Steamer Duck Lesser White-fronted Goose 58 136 Falkland Flightless Steamer Duck 138 Graylag Goose 60 Bar-headed Goose 63 Tribe Cairinini (Perching Ducks) 66 Spur-winged Goose 143 Ross Goose 69 144 Emperor Goose 72 White-winged Wood Duck 146

••• vii Page Page 148 Hottentot Teal 245 Hartlaub Duck 151 Garganey 247 Green 152 Blue-winged Teal 250 154 Cinnamon Teal 253 156 Red 256 158 Cape Shoveler 257 North American Wood Duck 160 260 Mandarin Duck 163 262 Australian Wood Duck 165 Pink-eared Duck 265 167 Marbled Teal 268

Tribe Merganettini () Tribe Aythyini (Pochards) Torrent Duck 173 Pink-headed Duck 273 Red-crested Pochard 274 Tribe Anatini Southern Pochard 277 (Dabbling or Surface-feeding Ducks) Rosybill 279 179 281 Salvadori Duck 180 Eurasian Pochard 284 African Black Duck 182 Redhead 287 Eurasian 184 Ring-necked Duck 290 American Wigeon 187 Australasian White-eye 292 Chiloe Wigeon 190 Siberian White-eye (Baer Pochard) 295 192 Ferruginous White-eye 297 Gadwall 195 Madagascan White-eye 299 Baikal Teal 198 Tufted Duck 300 Green-winged Teal 200 New Zealand Scaup 303 Speckled Teal 204 Greater Scaup 305 Cape Teal 206 Lesser Scaup 307 Madagascan Teal 208 Gray Teal 209 Tribe (Sea Ducks) Chestnut Teal 211 Eider (Common Eider) 313 Brown Teal 214 King Eider 315 216 Spectacled Eider 318 North American Black Duck 219 Steller Eider 321 Meller Duck 222 324 Yellow-billed Duck 223 325 Gray Duck 224 Long-tailed Duck (Oldsquaw) 328 Philippine Duck 227 Black Scoter 330 Bronze-winged Duck 228 Surf Scoter 333 231 White-winged Scoter 335 Pintail 233 Bufflehead 338 Brown Pintail 236 Barrow 340 White-cheeked Pintail 238 343 Red-billed Pintail 241 346 Silver Teal 243 348 viii ••• Page Page Brazilian Merganser 351 White-headed Duck 373 Red-breasted Merganser 352 Maccoa Duck 376 Chinese Merganser 356 Argentine Blue-billed Duck 378 Goosander (Common Merganser) 357 Australian Blue-billed Duck 380 Auckland Island Merganser 361 Musk Duck 382 Tribe (Stiff-tailed Ducks) Sources Cited Black-headed Duck Glossary and Vernacular Name Derivations Masked Duck Index

Illustrations

Figures

Page Page 1. Diagram of Evolutionary Relationships 2. Diagram of External Features and of the Tribes and Genera of Anatidae xviii Areas of Waterfowl 396

Maps

1. Distribution of the Magpie Goose 2 22. Distribution of the Graylag Goose 62 2. Distribution of the Spotted Whistling 23. Distribution of the Bar-headed Goose 65 Duck 6 24. Distribution of the Snow Goose 68 3. Distribution of the Plumed Whistling 25. Distribution of the Ross Goose 71 Duck 9 26. Distribution of the Emperor Goose 73 4. Distribution of the Fulvous Whistling 27. Distribution of the Canada Goose 78 Duck 11 28. Distribution of the Barnacle Goose 82 5. Distribution of the Wandering Whistling 29. Distribution of the Brant 85 Duck 14 30. Distribution of the Red-breasted Goose 88 6. Distribution of the Lesser Whistling Duck 16 31. Distribution of the Cape Barren Goose 92 7. Distribution of the White-faced 32. Distribution of the Freckled Duck 98 Whistling Duck 19 33. Distribution of the Blue-winged Goose 103 8. Distribution of the Black-bellied and 34. Distribution of the Andean Goose 105 Cuban Whistling Ducks 23 35. Distribution of the Magellan Goose 107 9. Distribution of the White-backed Duck 25 36. Distribution of the 109 10. Distribution of the 28 37. Distribution of the Ashy-headed 11. Distribution of the Black Swan 32 Sheldgoose 111 12. Distribution of the Black-necked Swan 34 38. Distribution of the Ruddy-headed 13. Distribution of the Trumpeter Swan 37 Sheldgoose 112 14. Distribution of the Whooper Swan 41 39. Distribution of the Orinoco Goose 114 15. Distribution of the Whistling Duck 43 40. Distribution of the Egyptian Goose 116 16. Distribution of the Bewick Swan 46 41. Distribution of the 117 17. Distribution of the Coscoroba Swan 49 42. Distribution of the Cape Shelduck 120 18. Distribution of the Swan Goose 51 43. Distribution of the Australian and New 19. Distribution of the Bean Goose 54 Zealand Shelducks 123 20. Distribution of the White-fronted Goose 57 44. Distribution of the Northern Shelduck 128 21. Distribution of the Lesser White-fronted 45. Distribution of the Radjah Shelduck 130 Goose 59 46. Distribution of the 134

••• xi Page Page 47. Distribution of the Magellanic and 82. Distribution of the White-cheeked Falkland Flightless Steamer Ducks 137 Pintail 240 48. Distribution of the Spur-winged Goose 143 83. Distribution of the Red-billed Pintail 242 49. Distribution of the Muscovy Duck 145 84. Distribution of the Silver Teal 245 50. Distribution of the White-winged Wood 85. Distribution of the Hottentot Teal 247 Duck 147 86. Distribution of the Blue-winged Teal and 5l. Distribution of the Comb Duck 150 Garganey 249 52. Distribution of the Hartlaub Duck 152 87. Distribution of the Cinnamon Teal 255 53. Distribution of the 153 88. Distribution of the Red Shoveler 256 54. Distribution of the Cotton Pygmy Goose 156 89. Distribution of the Cape Shoveler 258 55. Distribution of the African Pygmy 90. Distribution of the Australasian Goose 157 Shoveler 261 56. Distribution of the Ringed Teal 159 91. Distribution of the Northern Shoveler 264 57. Distribution of the North American 92. Distribution of the Pink-eared Duck 267 Wood Duck 162 93. Distribution of the Marbled Teal 269 58. Distribution of the Mandarin Duck 164 94. Distribution of the Red-crested Pochard 276 59. Distribution of the Australian Wood 95. Distribution of the Southern Pochard 278 Duck 166 96. Distribution of the Rosybill 280 60. Distribution of the Brazilian Teal 168 97. Distribution of the Canvasback 283 6l. Distribution of the Torrent Duck 172 98. Distribution of the Eurasian Pochard 285 62. Distribution of the Salvadori Duck and 99. Distribution of the Redhead 289 Blue Duck 178 100. Distribution of the Ring-necked Duck 291 63. Distribution of the African Black Duck 183 10l. Distribution of the Australasian 64. Distribution of the Eurasian Wigeon 186 White-eye 293 65. Distribution of the American Wigeon 189 102. Distribution of the Siberian White-eye 296 66. Distribution of the Chiloe Wigeon 191 103. Distribution of the Ferruginous 67. Distribution of the Falcated Duck 194 White-eye 298 68. Distribution of the Gadwall 196 104. Distribution of the Tufted Duck 302 69. Distribution of the Baikal Teal 199 105. Distribution of the Greater Scaup, 70. Distribution of the Green-winged Teal 202 New Zealand Scaup, and Madagascan White-eye 71. Distribution of the Speckled Teal 205 306 106. Distribution of the Lesser Scaup 72. Distribution of the Cape Teal and 309 Madagascan Teal 207 107. Distribution of the Eider 312 108. 73. Distribution of the Gray Teal 210 Distribution of the King Eider 317 74. Distribution of the Chestnut Teal and 109. Distribution of the Spectacled Eider 320 Brown Teal 213 110. Distribution of the Steller Eider 323 75. Distribution of the Mallard 218 111. Distribution of the Harlequin Duck 327 76. Distribution of the North American 112. Distribution of the Long-tailed Duck 329 Black Duck 221 113. Distribution of the Black Scoter 332 77. Distribution of the Gray, Philippine, 114. Distribution of the Surf Scoter 335 Yellow-billed, and Meller Ducks 226 115. Distribution of the White-winged Scoter 337 78. Distribution of the Bronze-winged Duck 230 116. Distribution of the Bufflehead 340 79. Distribution of the Crested Duck 232 117. Distribution of the Barrow Goldeneye 342 80. Distribution of the Pintail 235 118. Distribution of the Common Goldeneye 344 81. Distribution of the Brown Pintail 238 119. Distribution of the Hooded Merganser 347 xii ...... Page Page 120. Distribution of the Smew 349 127. Distribution of the Ruddy Duck 371 12l. Distribution of the Brazilian Merganser 352 128. Distribution of the White-headed Duck 375 122. Distribution of the Red-breasted 129. Distribution of the Maccoa Duck 377 Merganser 354 130. Distribution of the Argentine Blue-billed 123. Distribution of the Chinese Merganser 356 Duck 379 124. Distribution of the Goosander 359 13l. Distribution of the Australian Blue-billed 125. Distribution of the Black-headed Duck 366 Duck 381 126. Distribution of the Masked Duck 368 132. Distribution of the Musk Duck 384

Color Plates

following page 96 following page 176 l. White-faced Whistling Duck 29. Comb Duck 2. Magpie Goose 30. Spur-winged Goose 3. Spotted Whistling Duck 3l. Muscovy Duck 4. Plumed Whistling Duck 32. Hartlaub Duck 5. Lesser Whistling Duck 33. African Pygmy Goose 6. Black-bellied Whistling Duck 34. Ringed Teal 7. African White-backed Duck 35. Mandarin Duck 8. Mute Swan 36. Torrent Duck 9. Black Swan 37. American Wigeon 10. Black-necked Swan 38. Chiloe Wigeon 1I. Trumpeter Swan 39. Falcated Duck 12. Whooper Swan 40. Baikal Teal 13. Bewick Swan 4I. Cape Teal 14. Coscoroba Swan 42. Crested Duck 15. Lesser White-fronted Goose 43. Bronze-winged Duck 16. Graylag Goose 44. White-cheeked Pintail 17. Snow Goose 45. Garganey 18. Barnacle Goose 46. Red-crested Pochard 19. Brant 47. Eurasian Pochard 20. Hawaiian Goose 48. Redhead 21. Red-breasted Goose 49. Greater Scaup 22. Cereopsis Goose 50. American Eider 23. Andean Goose 5l. King Eider 24. Magellan Goose 52. Spectacled Eider 25. Ashy-headed Sheldgoose 53. Steller Eider 26. Egyptian Goose 54. Long-tailed Duck 27. Cape Shelduck 55. Harlequin Duck 28. Magellanic Flightless Steamer Duck 56. Bufflehead 57. Barrow Goldeneye 58. Hooded Merganser 59. White-headed Duck

• •• xiii

Preface

Of all the books that I have seen and wished that one can be found in Delacour's monograph, and likewise day I could personally own, perhaps the ones that to reduce descriptive behavioral information to a have most set me to daydreaming are the four-vol­ minimum whenever it had already been summarized ume set A Natural History of the Ducks, by John C. in my Handbook of Waterfowl Behavior. After con­ Phillips. My first contact with this magnificent siderable deliberation, it was decided to include monograph was at Cornell University, where it could descriptions as an aid to identification but be seen in the rare book collection. I spent countless not to attempt to describe all molts and . hours there during the late 1950s savoring the text The "Natural History" section of each species ac­ and plates. At about that same time, the first vol­ count was considered to be the nuclear element, and umes of Jean Delacour's The Waterfowl of the World an effort was made to include newer or at least dif­ were appearing, and these too gave me a chance to ferent information from that summarized by Dela­ dream about such entrancing birds as torrent ducks, cour. The "Status" and "Relationships" sections were steamer ducks, musk ducks, and magpie geese. Since added after the text was well underway, the former then it has been my good fortune to see all of these because of the increasing incidence of rare and en­ marvelous birds in life, and they have enriched my dangered forms among the waterfowl, and the latter own life enormously. The waterfowl family is so rich because more information on phyletic relationships is in ecological diversity, in evolutionary lessons, and available now than in 1965, when I last dealt com­ in behavioral complexities that I have never tired of prehensively with this problem in the Handbook of either reading or writing about it. Waterfowl Behavior. Only two of my previous books, Waterfowl: Their Except in a few instances, I have followed the taxo­ Biology and Natural History and the earlier Hand­ nomic treatment used in the Handbook, both as to book of Waterfowl Behavior, have dealt with the en­ major taxonomic categories and sequences of species tire waterfowl family in a comprehensive fashion, within such categories. The two exceptions constitute and neither attempted to provide a systematic review the recognition of special monotypic tribes for the of the biology of every species. Indeed, I have re­ Cape Barren goose and the torrent duck, since recent peatedly thought about undertaking such an effort studies on both species have cast doubt on their pre­ but each time have shrunk back at the thought of the sumed phyletic affinities and thus tribal recognition huge number of species to be considered and of the seemed to provide the best interim solution. The only vast literature that has accumulated on the group. other major change from the used in the Even in the 1920s it required four volumes for John Handbook is the transfer of the Thalassornis C. Phillips to review the biology of the waterfowl from the stifftails to the whistling duck tribe. This ac­ family exclusive of the swans and geese, and Jean tion was based on my own studies after the publica­ Delacour found that the same number of volumes tion of the Handbook and has received independent was needed for his treatment of the entire family. support from other investigators. A few changes I decided that, in spite of the concessions that have also been made in species limits and in sub­ would need to be made, a shorter approach to the species recognized, and a moderate number of sug­ family Anatidae was possible, provided that a maxi­ gested changes in English vernacular names have mum condensation of text be attained, primarily been adopted. In part these have been to avoid the through avoiding repetitous material such as separate use of "common" for vernacular names, but also to accounts for each subspecies. I also decided to ex­ allow for the consistent formation of distinctive ver­ clude taxonomic synonymies, extensive mensural nacular names for each subspecies by simply adding data, and most avicultural information, all of which an appropriate adjectival prefix. The group of four

••• xv swans known as the "northern swans" has proven Delacour for encouraging me to undertake the book refractory to such treatment, partly because of their in spite of its potential competition with his own questionable taxonomic status and their still uncer­ monograph. Not only have I relied extensively on the tain ancestral history. Thus, they have been given volumes by Delacour and Phillips but also on the separate accounts and treated as if they are "good" regional volumes by H. J. Frith (), G. species, in spite of the obvious fact that they are not. Dementiev and N. Gladkov (U.S.S.R.), P. A. Clan­ There are thus 148 separate species accounts in the cey (South ), S. Ali and D. Ripley (India), text. and C. Tso-hsin () for various mensural and The writing of the text was begun in the spring of descriptive information, sometimes without citations 1976 and continued through most of 1977, so that to these sources. I also wish to thank David Skead for later literature was included only if it was regarded as providing unpublished data on South African of critical importance. Further, the number of litera­ species. ture citations was held to an absolute minimum, but The distribution maps have been based on a va­ one or more "Suggested Readings" for each species riety of sources, including the volumes just men­ was included to provide additional access to the near­ tioned. The photographs are mostly of captive birds, ly endless literature on the waterfowl family. I would taken at the Wildfowl Trust. All the photographs and like to express my sincere appreciation to the School line drawings are my own, although several of the of Life Sciences for supporting this work and for al­ drawings are based on published photographs taken lowing me the time needed to complete it. by a variety of other photographers. In cases of rare In addition, I must again thank a number of in­ or extinct species I have sometimes used photographs dividuals for their invaluable aid. These include vir­ of live birds representing close relatives for propor­ tually all of the staff of the Wildfowl Trust, but tions and postures, but virtually none of the draw­ especially Sir Peter Scott, Dr. Geoffrey Matthews, ings lack a documentary base. and Dr. Janet Kear. I would also like to thank Jean

xvi ...... Introduction to the Family Anatidae

Inasmuch as the primary purpose of this book is to Tribe Tachyerini: Steamer ducks (1 genus, 3 provide information on each of the species of the species) waterfowl family in a standardized format and easily Tribe Cairinini: Perching ducks (9 genera, 13 accessible manner, it is important that the reader species) have some knowledge of the basis for my sequential Tribe Merganettini: Torrent duck (1 genus and organization of these species. A variety of attempts species) to provide a "natural" classification, or one that best Tribe Anatini: Dabbling or surface-feeding reflects actual evolutionary relationships, of the ducks (4 genera, 39 species) family Anatidae have been made in recent years, Tribe Aythyini: Pochards (3 genera, 16 species) with most of them being minor variations on a Tribe Mergini: Sea ducks (8 genera, 20 species) scheme first proposed by Jean Delacour and Ernst Tribe Oxyurini: Stiff-tailed ducks (3 genera, 8 Mayr in 1945. In this landmark classification, em­ species) phasis was given to the association of species at the tribal level, rather than to the fragmentation of the These groups are believed to be related to one family into a large number of subfamilies, as in another in the manner shown in figure 1, which in­ earlier classifications. dicates the probable relationships of the 13 tribes and My own behavioral studies of the family resulted 43 genera recognized in this book. Similar diagrams in a proposed classification for the group in 1961, showing species relationships for each of the major which was utilized in my subsequent books (1965a, tribes have been published earlier Oohnsgard, 1968a) and has been subject to only minor modifica­ 1961a), and with relatively few more recent modifi­ tion since that time based on new information from cations still provide the basis for the sequence in my own and other studies. In brief, the family which individual species are considered in this book. Anatidae is here regarded to be composed of 3 sub­ Behavioral and anatomical characteristics that pro­ families, 13 tribes, 43 genera, and 148 recent species vide the basis for the association of these species into as follows: tribes and subfamilies are those which are regarded as particularly significant, and thus it is worth review­ Family Anatidae (ducks, geese, and swans) ing such "emergent" characteristics before the spe­ Subfamily Anseranatinae cies-by-species consideration of the entire family. Tribe Anseranatini: Magpie goose (1 genus and species) Subfamily SUBFAMILY ANSERANATINAE Tribe Dendrocygnini: Whistling or tree ducks (2 Tribe Anseranatini (Magpie Goose) genera, 9 species) Tribe Anserini: Swans and true geese (4 genera, This subfamily is composed of a single or mono typic 21 species) tribe, genus, and species, the magpie goose, which Tribe Cereopsini: Cape Barren goose (1 genus differs so much from the remainder of the family that and species) some persons have argued that it might best be plac­ Tribe Stictonettini: Freckled duck (1 genus and ed in a separate family. However, it also provides species) such important transitional characteristics between Subfamily the more typical waterfowl and the South American Tribe Tadornini: Sheldgeese and Shelducks (5 of the family Anhimidae that it is perhaps genera, 15 species) most useful to retain it in the Anatidae as a "land-

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mark" form, against which the other species can use­ the species' semiterrestrial adaptations and its perch­ fully be measured. ing behavior. Like the other Anatidae, the magpie Foremost among the magpie goose's peculiarities goose molts its major flight (primaries and are its feet, which are only slightly webbed, and its secondaries) only once a year; but unlike virtually all unusually long hind toes, which are associated with of these other species, the magpie goose molts very xviii +++ gradually, with only a few feathers being absent at the sexes are essentially identical in plumage but anyone time, and thus the birds are continuously often differ somewhat in body weight and, occa­ able to fly. There is no flightless period of increased sionally, in voice. Although the whistling ducks have vulnerability to , and the greatly prolonged relatively long hind toes, not even they perch as well period of molting is not a major disadvantage to this as does the magpie goose, and the other species do tropically adapted and nearly sedentary species. not perch at all. All of the species lack iridescent A second unique anatomical feature of this species coloration, even on the wings. is the greatly elongated trachea in adults of both sexes, which in adult males may reach 150 centi­ Tribe Dendrocygnini (Whistling Ducks) meters. The magpie goose is unlike some swans in which the trachea is convoluted inside the sternum. The whistling ducks are a group of nine species, eight Rather, in this species it penetrates the area between of which are readily placed in the single genus Den­ the breast muscles and the skin, in a manner com­ drocygna, or "tree ducks." However, they are not parable to that in certain species of Oohns­ primarily perching birds, and thus "whistling ducks" gard, 1961b). As in the latter species, the resulting is a better vernacular designation than the frequently call is unusually low-pitched, and the elongated used "tree ducks." In nearly all species both sexes ut­ trachea probably functions as an effective resonator ter a clear, often multisyllabic whistle that readily of low-frequency sounds. Other than this specializa­ identifies the as to its species and apparently tion, however, the anatomy of the sound-producing provides important communication signals in these syrinx is relatively simple and gooselike. rather gregarious birds. Most of the species are tropi­ A third remarkable feature of the magpie goose is cal in distribution, and in all of them the pair bond related to its breeding behavior. The birds often forrr, seems to be permanent and potentially lifelong. Both a trio consisting of a male and two females that drop sexes share equally in brood-rearing responsibilities, their in a single nest, with all three birds sharing and in some species the male is known to participate the incubation responsibilities. This behavior, first in incubation as well. Unlike those of geese and observed in captive birds Oohnsgard, 1961c), was swans, the downy young are distinctively patterned. subsequently found to be typical also of wild in­ Although the downy young of the white-backed dividuals, and appears to be unique in the family. In duck differ somewhat from those of Dendrocygna, addition, magpie geese are the only waterfowl that the species shares a sufficiently large number of feed their young directly, the adults pickip..g up edible behavioral and anatomical traits to be tentatively materials in their bills and dropping them in front of included in this tribe. their downy young. This behavior has its counter­ part in the "tidbitting" behavior of many gal­ Tribe Anserini (Swans and True Geese) linaceous birds, but has not yet been reported from other Anatidae. The swans and true geese are moderately to extreme­ ly large waterfowl, which in common with the whis­ tling ducks have plumage patterns that are alike in SUBFAMILY ANSERINAE both sexes and lack iridescent coloration, and they also possess reticulated scale patterns on the tarsal The subfamily Anserinae includes the whistling surfaces. Most of the 20 species are found in the duck, swans, true geese, and two "aberrant" species. cooler parts of the , the excep­ Like the magpie goose, these species undergo only a tions being 3 species of Southern Hemisphere swans. single molt per year, which occurs after the breeding All are vegetarians, obtaining much of their food season and during which all feathers are replaced. from terrestrial surface vegetation in the case of However, the flight feathers are shed almost simul­ geese, and from subsurface aquatic vegetation in the taneously, so that the birds are unable to fly for a case of swans. The patterns of the downy young tend period of from about three to six weeks. In all of the to be pale and simple, without strong head or back included species the unfeathered part of the lower patterning, and in most species the adult plumages leg, or tarsus-like that of the magpie goose-has an are also fairly simple, with whites and blacks often irregular network, or reticulated pattern, of scales predominating. The white plumage of most adult throughout. Except in two semi terrestrial geese, how­ swans appears to be related to visibility needs associ­ ever, the front toes are fully webbed. In all species ated with their high degree of territorial spacing. The

...... xix most highly territorial swans are also the most distinct plumages, breeding (nuptial) and non­ strongly vocal ones, for the same reason. However, breeding (winter, or eclipse). In some species the vocalizations of the sexes are very similar, and usu­ breeding plumage of the male closely resembles that ally differ only in minor pitch characteristics. Most of the females, but more often the sexes have quite of the swans and geese are quite strongly migratory, different breeding plumages, in conjunction with but the insular Hawaiian goose not only has become more frequently disrupted and renewed pair bonds. nonmigratory but also has become semiterrestrial Likewise, the adult voices of the two sexes are nor­ and has evolved reduced toe webbing as it has mally very different in these species as the result of adapted to a mountainous existence on old lava sexual differences in the structure of the syrinx and fields. sometimes also the trachea. Unlike that in the pre­ vious subfamilies, the front surface of the lower tar­ sus has a linearly arranged (scutellated) scale pattern. Tribe Cereopsini (Cape Barren Goose) Iridescent coloration is frequently present in the Several recent studies have suggested that this un­ plumage, particularly among males, although on fe­ usual Australian gooselike bird is not very closely males it is usually restricted to the region of the related to either the true geese or the sheldgeese, secondaries of the wings. Such colorful patterns are although it shares some traits with both groups. It called speculums and are usually species-specific. has some unique traits, such as its unusually swollen In those species in which the sexes are quite dif­ bill, its adaptations to a terrestrial existence on the ferent, or dimorphic, in size and appearance, the coastline and islands of the Bass Strait, and its very male is typically larger, more brilliantly patterned, simple syringeal structure. Like the freckled duck, it and more aggressive. The plumages of juveniles and is perhaps best considered as a survivor of a group the nonbreeding plumages of males generally re­ transitional between the two major subfamilies of semble that of the adult female, and lack both irides­ waterfowl that exist at the present time. cent coloration on the body and the finely barred, or vermiculated, markings on the back, sides, and flanks that are typical of many of the species of this Tribe Stictonettini (Freckled Duck) subfamily. The patterns of the downy young are Even more than the Cape Barren goose, the freckled often quite contrasting and distinctive, and usually duck exhibits an assortment of anatomical traits that include spotting and striping on the head and back. suggest affinities with the geese and swans, in spite of In no cases are males known to assist in incubation, a fairly typical ducklike appearance and foraging but male participation in brood rearing is often behavior. Its unpatterned downy young, its remark­ variable within and between tribes, and is related ably primitive syringeal structure, and its reticulated primarily to such ecological factors as length and tarsus all strongly argue for the position that this regularity of the breeding seasons. species is the sole survivor of a very ancient water­ fowl lineage, with no near living relatives. A detailed Tribe Tadornini (Sheldgeese and Shelducks) study of its social behavior is greatly to be desired, for like the magpie goose, it provides an unequaled The most gooselike species of the subfamily Anatinae opportunity to gain insight into the evolutionary his­ are the sheldgeese, which together with the closely tory of the waterfowl group through observation of a related shelducks constitute the tribe Tadornini. This unique contemporary form. group of 14 species has a worldwide distribution ex­ cept for North America. The typical sheldgeese are grazing birds, the ecological counterparts of the true SUBFAMILY ANATINAE geese, while the shelducks are mostly wading and dabbling birds, frequently feeding to a large extent The third and largest subfamily, Anatinae, includes on aquatic . In all of the species the the sheldgeese, shelducks, and all of the typical adult male has an enlarged bony bulla in the syrinx, ducks. It is subdivided into a varying number of which is evidently responsible for the whistling or tribes by different authorities; nine are recognized whistlelike sounds that are important in sexual and here. In most if not all of the species included in this aggressive displays. The females of all species have subfamily there are two molts of the body feathers more reedy and typically gooselike or ducklike notes. per year, and consequently there are usually two In all species the males are larger than the females, xx +++ and often differ appreciably from them in plumage. long or irregular breeding seasons. Pair bonding is Although pair bonds are sometimes said to be perma­ generally seasonally established in the species that nent in the group, in some species at least they may breed in temperate climates, while in the more be ruptured through interactions with outside males, tropical and seemingly more primitive species the and a premium is placed on male aggressiveness dur­ pair bonds are weak and in a few cases apparently ing sexual competition for mates. The wings are used even nonexistent. The patterns of the downy young in fighting and in most species they are strongly pat­ are typically well marked and characterized by terned, with white coverts and iridescent coloration white or yellow spots and stripes on a darker on the secondaries or their greater coverts. The background; in many species they are not readily downy young are also typically patterned contrast­ separable from the downy young of dabbling ingly in the shelducks and sheldgeese, a characteristic ducks. The two tribes are obviously closely related, shared with various other cavity-nesting waterfowl. and taxonomists are not agreed on which group some species such as the ringed teal, pink-eared duck, and Brazilian teal should be included in. Tribe Tachyerini (Steamer Ducks) However, to merge the tribes would make a com­ The South American steamer ducks have at times paratively large tribe and tend to obscure the well­ been included with the shelducks, but differ enough defined differences in the behavior patterns and from them in structure and behavior to be regarded breeding biologies of these two groups. as a separate but closely related tribe. There are three species very similar in appearance; two are Tribe Merganettini (Torrent Duck) essentially flightless. All are found off the coasts of southern and the , This remarkable stream-dwelling duck of the An­ where they feed on mollusks and other marine in­ dean mountains is certainly one of the most vertebrates. The males differ slightly from females specialized of all waterfowl, and in part its in their plumage and vocalizations, but in both anatomical and behavioral specializations have sexes iridescent coloration is totally lacking and on­ obscured its basic relationships, which appear to be ly a simple white speculum is present on the wings. with either the perching ducks or the dabbling Like the shelducks, however, they are highly ag­ ducks. In the absence of definitive evidence, it gressive during social display activities. The downy seems reasonable to maintain a separate tribe for young to some extent resemble those of shelducks, this species, which exists as a series of relatively but have less spotting and less contrasting colora­ isolated populations between and Tierra tion. del Fuego that vary greatly in male plumage char­ acteristics. Torrent ducks are essentially cavity- or ledge-nesting forms, although only a few nests have Tribe Caidnini (Perching Ducks) been found, and much still remains to be learned of This group of 13 species of primarily perching their breeding biology. waterfowl is a rather heterogeneous assemblage of birds that are not easily characterized. In addition Tribe Anatini (Dabbling Ducks) to being generally perching and cavity-nesting, they also have fairly wide, rounded wings, elongated The dabbling, or surface-feeding, ducks are, to tails, and sometimes also relatively long legs. Many judge from the number and abundance of the in­ of the species exhibit a great deal of iridescent col­ cluded species, the most successful of all waterfowl. oration in their plumage, even among females, and This tribe includes all of the "puddle ducks" that in a few species nearly the entire body plumage is constitute most of the important game species iridescent. In some of these brilliantly colored throughout the world. Of the 39 species making up forms, such as the comb duck, muscovy duck, and the tribe, all but 3 can readily be placed in the the spur-winged goose, there is a great difference in single genus . The tribe has a worldwide the sizes of the sexes even though they may be very distribution, with some of the species occurring on similar in plumage patterning. In spite of their several continents and having extensive transcon­ brilliant plumages, males of only a few of the tinental migration patterns. However, most of the species exhibit definite eclipse plumages, probably species are temperate-breeding forms and are in part because most of the forms are tropical, with generally adapted to shallow, marshy habitats

••• xxi where food can be obtained from near the surface birds. They are found in fresh-water as well as by dabbling or tipping-up. In most species the marine habitats, and primarily have Northern males have fairly elaborate and colorful breeding Hemisphere distribution patterns. Except for the plumages, whereas the ground-nesting females are two isolated Southern Hemisphere species, all of mostly cryptically patterned with buff and brown. the sea ducks have considerable sexual dimorphism In nearly all species both sexes have iridescent wing in plumage. Males in breeding plumage are usually speculum markings, and in all species the trachea elaborately patterned, often with predominantly has an enlarged bulla at the syrinx. Pair bonds are black and white markings, which probably serve generally reformed each year during the non­ very well to localize and identify these birds at breeding season, but in some species with long considerable distances in their marine environ­ breeding seasons the pair bonds are relatively per­ ments. However, iridescent coloration is generally manent and the males remain with their offspring limited to the head, and the wing speculum pat­ and help to protect them. , instead of being iridescent, are generally white and black. Nesting is usually done on fairly open shoreline or in grassy tundra, as in the eiders Tribe Aythyini (Pochards) and long-tailed duck, or in cavities, under heavy This tribe of primarily fresh-water diving ducks brush, or in similar well-concealed locations, as in contains 16 species that collectively may be called the goldeneyes, mergansers, and harlequin duck. pochards. The tribe has a nearly worldwide distri­ Females of the ground-nesting forms approach the bution, but only a few of its species have ranges patterns found among female dabbling ducks in that extend beyond a single continent. In addition their cryptic coloration, and the downy young of to 15 species that are very similar in body propor­ these species are also rather brownish and obscure­ tions and diving adaptations, the apparently ex­ ly patterned. Females of the hole-nesting species are tinct pink-headed duck has anatomical character­ more uniformly brownish and their ducklings are istics that approach those of the pochards and it frequently contrastingly spotted with white and clearly should be included in this tribe. All of the dark markings. typical pochards are adept at diving and possess large feet, with long outer toes and strongly lobed Tribe Oxyurini (Stiff-tailed Ducks) hind toes. Their legs are placed quite far apart and are situated farther back on the body than in other The stiff-tailed ducks are characterized by several ducks, making the birds relatively awkward on unusual features in addition to their long, stiffened land but improving their diving efficiency. They tail feathers that serve as underwater rudders. also have a heavier body size to wing-surface ratio, Among other adaptations are their very large feet, forcing them to run for some distance over the which are placed so far back on the body that it is water when taking flight, in contrast to the "spring­ difficult for these birds to walk on land. The body ing" takeoff of dabbling ducks. Although sexual feathers are small, numerous, and have a grebelike dimorphism occurs to some extent in all species, sheen, and the wings are so short that takeoff and male plumage patterns are generally not especially flight is attained with difficulty. All of the species complex, and the wing speculum patterns are either have relatively short and thick necks, which in lacking or limited to gray or white stripes. The males can be enlarged by the inflation of various females of pochards are usually rather uniformly internal structures during sexual display. Although brownish, and lack the very distinct patterning of all of the species exhibit some dimorphism of size female dabbling ducks. Nesting is usually done on or coloration, the plumage patterns are mostly land near water or, more commonly, in beds of shades of gray, ruddy brown, and black, with con­ emergent vegetation. The downy young tend to be trasting white markings largely limited to the head. weakly or obscurely patterned with shades of Only one species, the masked duck, exhibits a dis­ yellow and dark brown. tinct speculum, and this species is furthermore the only one that is able to attain flight easily. The downy plumage patterns are usually very similar to Tribe Mergini (Sea Ducks) those of the adult females, which tend to be incon­ The sea duck tribe, which here includes the eiders, spicuously patterned with grays and browns. Non­ consists of 20 species that are all superb diving breeding plumages of males also closely resemble

xxii ••• those of the females, and during this period the more closely approaches the dabbling ducks. In ad­ ruddy coloration of the typical forms is largely dition to providing an apparent evolutionary link lost, as is the unusual blue color of the male's bill. with that group, the black-headed duck is also the Two species that do not fit this general pattern are only species of Anatidae that seems to have be­ the Australian musk duck and the South American come a total social parasite, abandoning its nest­ black-headed duck. The musk duck may readily be building tendencies in favor of dropping its eggs in considered a typical stiff tail that, under evolu­ the nests of a variety of host species. In this sense tionary pressures associated with a highly com­ it is one of the most specialized of all waterfowl petitive mating system favoring strength and species, although its parasitic adaptations are aggressiveness, has become remarkably large and relatively primitive by comparison with those of has evolved bizarre male displays. However, the some of the better-known species of social para­ black-headed duck lacks many of the traits of the sites. stiff-tailed ducks, and in many anatomical features

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