Grzimek Animal Life Encyclopedia Volume 11 Birds IV ( PDFDrive )

Grzimek’s

Animal Life Encyclopedia

Second Edition

●●●●

Grzimek’s

Animal Life Encyclopedia

Second Edition

●●●●

Volume 11
Birds IV

Jerome A. Jackson, Advisory Editor
Walter J. Bock, Taxonomic Editor

Donna Olendorf, Project Editor
Joseph E. Trumpey, Chief Scientific Illustrator

Michael Hutchins, Series Editor

In association with the American Zoo and Aquarium Association

Grzimek’s Animal Life Encyclopedia, Second Edition

Volume 11: Birds IV

Project Editor Permissions Product Design
Donna Olendorf Kim Davis Tracey Rowens, Jennifer Wahi

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Kretschmann, Melissa C. McDade, Mark
Springer

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LIBRARY OF CONGRESS CATALOGING-IN-PUBLICATION DATA

Grzimek, Bernhard.
[Tierleben. English]
Grzimek’s animal life encyclopedia.— 2nd ed.
v. cm.

Includes bibliographical references.
Contents: v. 1. Lower metazoans and lesser deuterosomes / Neil Schlager, editor
— v. 2. Protostomes / Neil Schlager, editor — v. 3. Insects / Neil Schlager, editor —
v. 4-5. Fishes I-II / Neil Schlager, editor — v. 6. Amphibians / Neil Schlager, editor
— v. 7. Reptiles / Neil Schlager, editor — v. 8-11. Birds I-IV / Donna Olendorf, edi-
tor — v. 12-16. Mammals I-V / Melissa C. McDade, editor — v. 17. Cumulative
index / Melissa C. McDade, editor.
ISBN 0-7876-5362-4 (set hardcover : alk. paper)
1. Zoology—Encyclopedias. I. Title: Animal life encyclopedia. II.
Schlager, Neil, 1966- III. Olendorf, Donna IV. McDade, Melissa C. V. American Zoo
and Aquarium Association. VI. Title.
QL7 .G7813 2004

590Ј.3—dc21
2002003351

Printed in the United States of America
10 9 8 7 6 5 4 3 2 1

Recommended citation: Grzimek’s Animal Life Encyclopedia, 2nd edition. Volumes 8–11, Birds I–IV, edited by Michael
Hutchins, Jerome A. Jackson, Walter J. Bock, and Donna Olendorf. Farmington Hills, MI: Gale Group, 2002.

•••••

Contents

Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . viii Family: Tropicbirds . . . . . . . . . . . . . . . . . . . . . . . . . . 187
How to use this book . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi Family: Frigatebirds . . . . . . . . . . . . . . . . . . . . . . . . . . 193
Advisory boards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii Family: Cormorants and anhingas . . . . . . . . . . . . . . 201
Contributing writers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xv Family: Boobies and gannets . . . . . . . . . . . . . . . . . . 211
Contributing illustrators . . . . . . . . . . . . . . . . . . . . . . . . . xviii Family: Pelicans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225

Volume 8: Birds I 3 Order CICONIIFORMES
What is a bird? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Herons, storks, spoonbills, ibis, and New World
Birds and humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 vultures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
Avian migration and navigation . . . . . . . . . . . . . . . . . . . . 37
Avian song . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Family: Herons and bitterns . . . . . . . . . . . . . . . . . . . 239
Avian flight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family: Hammerheads . . . . . . . . . . . . . . . . . . . . . . . 261
Family: Storks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265
Order STRUTHIONIFORMES Family: New World vultures . . . . . . . . . . . . . . . . . . 275
Tinamous and ratites . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 Family: Shoebills . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287
Family: Ibises and spoonbills . . . . . . . . . . . . . . . . . . 291
Family: Tinamous . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Family: Rheas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 Order PHOENICOPTERIFORMES
Family: Cassowaries . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Flamingos
Family: Emus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Family: Kiwis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 Family: Flamingos . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
Family: Moas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Family: Ostriches . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 Order FALCONIFORMES
Family: Elephant birds . . . . . . . . . . . . . . . . . . . . . . . 103 Diurnal birds of prey . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313

Order PROCELLARIIFORMES Family: Hawks and eagles . . . . . . . . . . . . . . . . . . . . . 317
Tubenosed seabirds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 Family: Secretary birds . . . . . . . . . . . . . . . . . . . . . . . 343
Family: Falcons and caracaras . . . . . . . . . . . . . . . . . 347
Family: Albatrosses . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Family: Shearwaters, petrels, and fulmars . . . . . . . . 123 Order ANSERIFORMES
Family: Storm-petrels . . . . . . . . . . . . . . . . . . . . . . . . 135 Ducks, geese, swans, and screamers . . . . . . . . . . . . . . . . 363
Family: Diving-petrels . . . . . . . . . . . . . . . . . . . . . . . . 143
Family: Ducks, geese, and swans . . . . . . . . . . . . . . . 369
Order SPHENISCIFORMES Family: Screamers . . . . . . . . . . . . . . . . . . . . . . . . . . . 393
Penguins
Order GALLIFORMES
Family: Penguins . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147 Chicken-like birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399

Order GAVIIFORMES Family: Moundbuilders . . . . . . . . . . . . . . . . . . . . . . . 403
Loons Family: Curassows, guans, and chachalacas . . . . . . . 413
Family: Guineafowl . . . . . . . . . . . . . . . . . . . . . . . . . . 425
Family: Loons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159 Family: Fowls and pheasants . . . . . . . . . . . . . . . . . . 433
Family: New World quails . . . . . . . . . . . . . . . . . . . . 455
Order PODICIPEDIFORMES
Grebes Order OPISTHOCOMIFORMES
Hoatzins
Family: Grebes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Family: Hoatzins . . . . . . . . . . . . . . . . . . . . . . . . . . . . 465
Order PELECANIFORMES
Pelicans and cormorants . . . . . . . . . . . . . . . . . . . . . . . . . 183 v

Grzimek’s Animal Life Encyclopedia

Contents

For further reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 469 Order STRIGIFORMES
Organizations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474 Owls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331
Contributors to the first edition. . . . . . . . . . . . . . . . . . . . 479
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 486 Family: Barn owls . . . . . . . . . . . . . . . . . . . . . . . . . . . 335
Aves species list . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 497 Family: Owls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
Geologic time scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 560
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 561 Order CAPRIMULGIFORMES
Nightjars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 367
Volume 9: Birds II 1
Order GRUIFORMES 5 Family: Oilbirds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 373
Cranes, rails, and relatives . . . . . . . . . . . . . . . . . . . . . . . . 11 Family: Frogmouths . . . . . . . . . . . . . . . . . . . . . . . . . 377
23 Family: Owlet-nightjars . . . . . . . . . . . . . . . . . . . . . . 387
Family: Mesites and roatelos . . . . . . . . . . . . . . . . . . 37 Family: Potoos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395
Family: Buttonquails . . . . . . . . . . . . . . . . . . . . . . . . . 41 Family: Nightjars . . . . . . . . . . . . . . . . . . . . . . . . . . . . 401
Family: Cranes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Family: Limpkins . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 Order APODIFORMES
Family: Kagus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 Swifts and hummingbirds . . . . . . . . . . . . . . . . . . . . . . . . 415
Family: Rails, coots, and moorhens . . . . . . . . . . . . . 77
Family: Sungrebes . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 Family: Swifts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 421
Family: Sunbitterns . . . . . . . . . . . . . . . . . . . . . . . . . . 91 Family: Tree swifts . . . . . . . . . . . . . . . . . . . . . . . . . . 433
Family: Trumpeters . . . . . . . . . . . . . . . . . . . . . . . . . . Family: Hummingbirds . . . . . . . . . . . . . . . . . . . . . . . 437
Family: Seriemas . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family: Bustards . . . . . . . . . . . . . . . . . . . . . . . . . . . . Order COLIIFORMES
Mousebirds
Order CHARADRIIFORMES
Gulls, terns, plovers, and other shorebirds . . . . . . . . . . . 101 Family: Mousebirds . . . . . . . . . . . . . . . . . . . . . . . . . . 469

Family: Jacanas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 Order TROGONIFORMES
Family: Painted snipes . . . . . . . . . . . . . . . . . . . . . . . . 115 Trogons
Family: Crab plovers . . . . . . . . . . . . . . . . . . . . . . . . . 121
Family: Oystercatchers . . . . . . . . . . . . . . . . . . . . . . . 125 Family: Trogons . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477
Family: Stilts and avocets . . . . . . . . . . . . . . . . . . . . . 133
Family: Thick-knees . . . . . . . . . . . . . . . . . . . . . . . . . 143 For further reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487
Family: Pratincoles and coursers . . . . . . . . . . . . . . . 151 Organizations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 492
Family: Plovers and lapwings . . . . . . . . . . . . . . . . . . 161 Contributors to the first edition. . . . . . . . . . . . . . . . . . . . 497
Family: Sandpipers . . . . . . . . . . . . . . . . . . . . . . . . . . 175 Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 504
Family: Seedsnipes . . . . . . . . . . . . . . . . . . . . . . . . . . . 189 Aves species list . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 515
Family: Sheathbills . . . . . . . . . . . . . . . . . . . . . . . . . . 197 Geologic time scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 578
Family: Gulls and terns . . . . . . . . . . . . . . . . . . . . . . . 203 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 579
Family: Auks, puffins, and murres . . . . . . . . . . . . . . 219
Volume 10: Birds III 1
Order PTEROCLIFORMES Order CORACIIFORMES 5
Sandgrouse Kingfishers, todies, hoopoes, and relatives . . . . . . . . . . . 25
31
Family: Sandgrouse . . . . . . . . . . . . . . . . . . . . . . . . . . 231 Family: Kingfishers . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Family: Todies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Order COLUMBIFORMES Family: Motmots . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Pigeons, doves, and dodos . . . . . . . . . . . . . . . . . . . . . . . . 241 Family: Bee-eaters . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Family: Rollers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Family: Pigeons and doves . . . . . . . . . . . . . . . . . . . . 247 Family: Hoopoes . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Family: Dodos and solitaires . . . . . . . . . . . . . . . . . . 269 Family: Woodhoopoes . . . . . . . . . . . . . . . . . . . . . . .
Family: Hornbills . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Order PSITTACIFORMES Order PICIFORMES
Parrots Woodpeckers and relatives . . . . . . . . . . . . . . . . . . . . . . . 85

Family: Parrots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275 Family: Jacamars . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Family: Puffbirds . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Order MUSOPHAGIFORMES Family: Barbets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Turacos and plantain eaters Family: Toucans . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Family: Honeyguides . . . . . . . . . . . . . . . . . . . . . . . . . 137
Family: Turacos and plantain eaters . . . . . . . . . . . . 299 Family: Woodpeckers, wrynecks, and piculets . . . . 147

Order CUCULIFORMES Order PASSERIFORMES
Cuckoos, anis, and roadrunners Perching birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169

Family: Cuckoos, anis, and roadrunners . . . . . . . . . 311 Family: Broadbills . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Family: False sunbirds and asities . . . . . . . . . . . . . . 187

vi Grzimek’s Animal Life Encyclopedia

Family: Pittas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193 Contents
Family: New Zealand wrens . . . . . . . . . . . . . . . . . . . 203
Family: Ovenbirds . . . . . . . . . . . . . . . . . . . . . . . . . . . 209 Family: Quail thrushes and whipbirds . . . . . . . . . . . 75
Family: Woodcreepers . . . . . . . . . . . . . . . . . . . . . . . 229 Family: Fantails . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Family: Ant thrushes . . . . . . . . . . . . . . . . . . . . . . . . . 239 Family: Monarch flycatchers . . . . . . . . . . . . . . . . . . 97
Family: Tapaculos . . . . . . . . . . . . . . . . . . . . . . . . . . . 257 Family: Australian robins . . . . . . . . . . . . . . . . . . . . . 105
Family: Tyrant flycatchers . . . . . . . . . . . . . . . . . . . . 269 Family: Whistlers . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
Family: Sharpbills . . . . . . . . . . . . . . . . . . . . . . . . . . . 291 Family: Pseudo babblers . . . . . . . . . . . . . . . . . . . . . . 127
Family: Manakins . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295 Family: Australian creepers . . . . . . . . . . . . . . . . . . . . 133
Family: Cotingas . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305 Family: Long-tailed titmice . . . . . . . . . . . . . . . . . . . 141
Family: Plantcutters . . . . . . . . . . . . . . . . . . . . . . . . . . 325 Family: Penduline titmice . . . . . . . . . . . . . . . . . . . . . 147
Family: Lyrebirds . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329 Family: Titmice and chickadees . . . . . . . . . . . . . . . . 155
Family: Scrub-birds . . . . . . . . . . . . . . . . . . . . . . . . . . 337 Family: Nuthatches and wall creepers . . . . . . . . . . . 167
Family: Larks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341 Family: Treecreepers . . . . . . . . . . . . . . . . . . . . . . . . . 177
Family: Swallows . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357 Family: Philippine creepers . . . . . . . . . . . . . . . . . . . 183
Family: Pipits and wagtails . . . . . . . . . . . . . . . . . . . . 371 Family: Flowerpeckers . . . . . . . . . . . . . . . . . . . . . . . . 189
Family: Cuckoo-shrikes . . . . . . . . . . . . . . . . . . . . . . . 385 Family: Pardalotes . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
Family: Bulbuls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395 Family: Sunbirds . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Family: Fairy bluebirds and leafbirds . . . . . . . . . . . . 415 Family: White-eyes . . . . . . . . . . . . . . . . . . . . . . . . . . 227
Family: Shrikes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 425 Family: Australian honeyeaters . . . . . . . . . . . . . . . . . 235
Family: Vanga shrikes . . . . . . . . . . . . . . . . . . . . . . . . 439 Family: Vireos and peppershrikes . . . . . . . . . . . . . . 255
Family: Waxwings and silky flycatchers . . . . . . . . . 447 Family: New World finches . . . . . . . . . . . . . . . . . . . 263
Family: Palmchats . . . . . . . . . . . . . . . . . . . . . . . . . . . 455 Family: New World warblers . . . . . . . . . . . . . . . . . . 285
Family: Hedge sparrows . . . . . . . . . . . . . . . . . . . . . . 459 Family: New World blackbirds and orioles . . . . . . 301
Family: Thrashers and mockingbirds . . . . . . . . . . . . 465 Family: Finches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323
Family: Dippers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475 Family: Hawaiian honeycreepers . . . . . . . . . . . . . . . 341
Family: Thrushes and chats . . . . . . . . . . . . . . . . . . . 483 Family: Waxbills and grassfinches . . . . . . . . . . . . . . 353
Family: Babblers . . . . . . . . . . . . . . . . . . . . . . . . . . . . 505 Family: Weavers . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
Family: Wrens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 525 Family: Sparrows . . . . . . . . . . . . . . . . . . . . . . . . . . . . 397
Family: Starlings and mynas . . . . . . . . . . . . . . . . . . . 407
For further reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 539 Family: Old World orioles and figbirds . . . . . . . . . 427
Organizations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 544 Family: Drongos . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
Contributors to the first edition. . . . . . . . . . . . . . . . . . . . 549 Family: New Zealand wattle birds . . . . . . . . . . . . . . 447
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 556 Family: Mudnest builders . . . . . . . . . . . . . . . . . . . . . 453
Aves species list . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 567 Family: Woodswallows . . . . . . . . . . . . . . . . . . . . . . . 459
Geologic time scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 630 Family: Magpie-shrikes . . . . . . . . . . . . . . . . . . . . . . . 467
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 631 Family: Bowerbirds . . . . . . . . . . . . . . . . . . . . . . . . . . 477
Family: Birds of paradise . . . . . . . . . . . . . . . . . . . . . . 489
Volume 11: Birds IV 1 Family: Crows and jays . . . . . . . . . . . . . . . . . . . . . . . 503
Family: Old World warblers . . . . . . . . . . . . . . . . . . . 25
Family: Old World flycatchers . . . . . . . . . . . . . . . . . 45 For further reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 525
Family: Australian fairy-wrens . . . . . . . . . . . . . . . . . 55 Organizations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 530
Family: Australian warblers . . . . . . . . . . . . . . . . . . . . 65 Contributors to the first edition. . . . . . . . . . . . . . . . . . . . 535
Family: Australian chats . . . . . . . . . . . . . . . . . . . . . . 69 Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 542
Family: Logrunners and chowchillas . . . . . . . . . . . . Aves species list . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 553
Geologic time scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 616
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 617

Grzimek’s Animal Life Encyclopedia vii

•••••

Foreword

Earth is teeming with life. No one knows exactly how many American Insects and searched through the section on moths
distinct organisms inhabit our planet, but more than 5 mil- and butterflies. It was a luna moth! My heart was pounding
lion different species of animals and plants could exist, rang- with the excitement of new knowledge as I ran to share the
ing from microscopic algae and bacteria to gigantic elephants, discovery with my parents.
redwood trees and blue whales. Yet, throughout this won-
derful tapestry of living creatures, there runs a single thread: I consider myself very fortunate to have made a living as
Deoxyribonucleic acid or DNA. The existence of DNA, an a professional biologist and conservationist for the past 20
elegant, twisted organic molecule that is the building block years. I’ve traveled to over 30 countries and six continents to
of all life, is perhaps the best evidence that all living organ- study and photograph wildlife or to attend related conferences
isms on this planet share a common ancestry. Our ancient and meetings. Yet, each time I encounter a new and unusual
connection to the living world may drive our curiosity, and animal or habitat my heart still races with the same excite-
perhaps also explain our seemingly insatiable desire for in- ment of my youth. If this is biophilia, then I certainly possess
formation about animals and nature. Noted zoologist, E.O. it, and it is my hope that others will experience it too. I am
Wilson, recently coined the term “biophilia” to describe this therefore extremely proud to have served as the series editor
phenomenon. The term is derived from the Greek bios mean- for the Gale Group’s rewrite of Grzimek’s Animal Life Ency-
ing “life” and philos meaning “love.” Wilson argues that we clopedia, one of the best known and widely used reference
are human because of our innate affinity to and interest in the works on the animal world. Grzimek’s is a celebration of an-
other organisms with which we share our planet. They are, imals, a snapshot of our current knowledge of the Earth’s in-
as he says, “the matrix in which the human mind originated credible range of biological diversity. Although many other
and is permanently rooted.” To put it simply and metaphor- animal encyclopedias exist, Grzimek’s Animal Life Encyclopedia
ically, our love for nature flows in our blood and is deeply en- remains unparalleled in its size and in the breadth of topics
grained in both our psyche and cultural traditions. and organisms it covers.

Our own personal awakenings to the natural world are as The revision of these volumes could not come at a more
diverse as humanity itself. I spent my early childhood in rural opportune time. In fact, there is a desperate need for a deeper
Iowa where nature was an integral part of my life. My father understanding and appreciation of our natural world. Many
and I spent many hours collecting, identifying and studying species are classified as threatened or endangered, and the sit-
local insects, amphibians and reptiles. These experiences had uation is expected to get much worse before it gets better.
a significant impact on my early intellectual and even spiri- Species extinction has always been part of the evolutionary
tual development. One event I can recall most vividly. I had history of life; some organisms adapt to changing circum-
collected a cocoon in a field near my home in early spring. stances and some do not. However, the current rate of species
The large, silky capsule was attached to a stick. I brought the loss is now estimated to be 1,000–10,000 times the normal
cocoon back to my room and placed it in a jar on top of my “background” rate of extinction since life began on Earth
dresser. I remember waking one morning and, there, perched some 4 billion years ago. The primary factor responsible for
on the tip of the stick was a large moth, slowly moving its this decline in biological diversity is the exponential growth
delicate, light green wings in the early morning sunlight. It of human populations, combined with peoples’ unsustainable
took my breath away. To my inexperienced eyes, it was one appetite for natural resources, such as land, water, minerals,
of the most beautiful things I had ever seen. I knew it was a oil, and timber. The world’s human population now exceeds
moth, but did not know which species. Upon closer exami- 6 billion, and even though the average birth rate has begun
nation, I noticed two moon-like markings on the wings and to decline, most demographers believe that the global human
also noted that the wings had long “tails”, much like the ubiq- population will reach 8–10 billion in the next 50 years. Much
uitous tiger swallow-tail butterflies that visited the lilac bush of this projected growth will occur in developing countries in
in our backyard. Not wanting to suffer my ignorance any Central and South America, Asia and Africa—regions that are
longer, I reached immediately for my Golden Guide to North rich in unique biological diversity.

viii Grzimek’s Animal Life Encyclopedia

Finding solutions to conservation challenges will not be Foreword
easy in today’s human-dominated world. A growing number
of people live in urban settings and are becoming increasingly a system of protected areas where wildlife can roam free from
isolated from nature. They “hunt” in super markets and malls, exploitation of any kind.
live in apartments and houses, spend their time watching tele-
vision and searching the World Wide Web. Children and While it is clear that modern conservation must take the
adults must be taught to value biological diversity and the needs of both wildlife and people into consideration, what will
habitats that support it. Education is of prime importance now the quality of human life be if the collective impact of short-
while we still have time to respond to the impending crisis. term economic decisions is allowed to drive wildlife popula-
There still exist in many parts of the world large numbers of tions into irreversible extinction? Many rural populations
biological “hotspots”—places that are relatively unaffected by living in areas of high biodiversity are dependent on wild an-
humans and which still contain a rich store of their original imals as their major source of protein. In addition, wildlife
animal and plant life. These living repositories, along with se- tourism is the primary source of foreign currency in many de-
lected populations of animals and plants held in profession- veloping countries and is critical to their financial and social
ally managed zoos, aquariums and botanical gardens, could stability. When this source of protein and income is gone,
provide the basis for restoring the planet’s biological wealth what will become of the local people? The loss of species is
and ecological health. This encyclopedia and the collective not only a conservation disaster; it also has the potential to
knowledge it represents can assist in educating people about be a human tragedy of immense proportions. Protected ar-
animals and their ecological and cultural significance. Perhaps eas, such as national parks, and regulated hunting in areas out-
it will also assist others in making deeper connections to na- side of parks are the only solutions. What critics do not realize
ture and spreading biophilia. Information on the conserva- is that the fate of wildlife and people in developing countries
tion status, threats and efforts to preserve various species have is closely intertwined. Forests and savannas emptied of wildlife
been integrated into this revision. We have also included in- will result in hungry, desperate people, and will, in the long-
formation on the cultural significance of animals, including term lead to extreme poverty and social instability. Dr.
their roles in art and religion. Grzimek’s early contributions to conservation should be rec-
ognized, not only as benefiting wildlife, but as benefiting lo-
It was over 30 years ago that Dr. Bernhard Grzimek, then cal people as well.
director of the Frankfurt Zoo in Frankfurt, Germany, edited
the first edition of Grzimek’s Animal Life Encyclopedia. Dr. Dr. Grzimek’s hope in publishing his Animal Life Encyclo-
Grzimek was among the world’s best known zoo directors and pedia was that it would “...disseminate knowledge of the ani-
conservationists. He was a prolific author, publishing nine mals and love for them”, so that future generations would
books. Among his contributions were: Serengeti Shall Not Die, “...have an opportunity to live together with the great diver-
Rhinos Belong to Everybody and He and I and the Elephants. Dr. sity of these magnificent creatures.” As stated above, our goals
Grzimek’s career was remarkable. He was one of the first in producing this updated and revised edition are similar.
modern zoo or aquarium directors to understand the impor- However, our challenges in producing this encyclopedia were
tance of zoo involvement in in situ conservation, that is, of more formidable. The volume of knowledge to be summa-
their role in preserving wildlife in nature. During his tenure, rized is certainly much greater in the twenty-first century than
Frankfurt Zoo became one of the leading western advocates it was in the 1970’s and 80’s. Scientists, both professional and
and supporters of wildlife conservation in East Africa. Dr. amateur, have learned and published a great deal about the
Grzimek served as a Trustee of the National Parks Board of animal kingdom in the past three decades, and our under-
Uganda and Tanzania and assisted in the development of sev- standing of biological and ecological theory has also pro-
eral protected areas. The film he made with his son Michael, gressed. Perhaps our greatest hurdle in producing this revision
Serengeti Shall Not Die, won the 1959 Oscar for best docu- was to include the new information, while at the same time
mentary. retaining some of the characteristics that have made Grzimek’s
Animal Life Encyclopedia so popular. We have therefore strived
Professor Grzimek has recently been criticized by some to retain the series’ narrative style, while giving the informa-
for his failure to consider the human element in wildlife con- tion more organizational structure. Unlike the original
servation. He once wrote: “A national park must remain a pri- Grzimek’s, this updated version organizes information under
mordial wilderness to be effective. No men, not even native specific topic areas, such as reproduction, behavior, ecology
ones, should live inside its borders.” Such ideas, although con- and so forth. In addition, the basic organizational structure is
sidered politically incorrect by many, may in retrospect actu- generally consistent from one volume to the next, regardless
ally prove to be true. Human populations throughout Africa of the animal groups covered. This should make it easier for
continue to grow exponentially, forcing wildlife into small is- users to locate information more quickly and efficiently. Like
lands of natural habitat surrounded by a sea of humanity. The the original Grzimek’s, we have done our best to avoid any
illegal commercial bushmeat trade—the hunting of endan- overly technical language that would make the work difficult
gered wild animals for large scale human consumption—is to understand by non-biologists. When certain technical ex-
pushing many species, including our closest relatives, the go- pressions were necessary, we have included explanations or
rillas, bonobos, and chimpanzees, to the brink of extinction. clarifications.
The trade is driven by widespread poverty and lack of eco-
nomic alternatives. In order for some species to survive it will Considering the vast array of knowledge that such a work
be necessary, as Grzimek suggested, to establish and enforce represents, it would be impossible for any one zoologist to
have completed these volumes. We have therefore sought spe-
Grzimek’s Animal Life Encyclopedia cialists from various disciplines to write the sections with

ix

Foreword detailed a treatment as did the birds and mammals. Due to
practical and financial considerations, the publishers could
which they are most familiar. As with the original Grzimek’s, provide only so much space for each animal group. In such
we have engaged the best scholars available to serve as topic cases, it was impossible to provide more than a broad overview
editors, writers, and consultants. There were some complaints and to feature a few selected examples for the purposes of il-
about inaccuracies in the original English version that may lustration. To help compensate, we have provided a few key
have been due to mistakes or misinterpretation during the bibliographic references in each section to aid those inter-
complicated translation process. However, unlike the origi- ested in learning more. This is a common limitation in all ref-
nal Grzimek’s, which was translated from German, this revi- erence works, but Grzimek’s Encyclopedia of Animal Life is still
sion has been completely re-written by English-speaking the most comprehensive work of its kind.
scientists. This work was truly a cooperative endeavor, and I
thank all of those dedicated individuals who have written, I am indebted to the Gale Group, Inc. and Senior Editor
edited, consulted, drawn, photographed, or contributed to its Donna Olendorf for selecting me as Series Editor for this
production in any way. The names of the topic editors, au- project. It was an honor to follow in the footsteps of Dr.
thors, and illustrators are presented in the list of contributors Grzimek and to play a key role in the revision that still bears
in each individual volume. his name. Grzimek’s Animal Life Encyclopedia is being published
by the Gale Group, Inc. in affiliation with my employer, the
The overall structure of this reference work is based on the American Zoo and Aquarium Association (AZA), and I would
classification of animals into naturally related groups, a disci- like to thank AZA Executive Director, Sydney J. Butler;
pline known as taxonomy or biosystematics. Taxonomy is the AZA Past-President Ted Beattie (John G. Shedd Aquarium,
science through which various organisms are discovered, iden- Chicago, IL); and current AZA President, John Lewis (John
tified, described, named, classified and catalogued. It should be Ball Zoological Garden, Grand Rapids, MI), for approving
noted that in preparing this volume we adopted what might be my participation. I would also like to thank AZA Conserva-
termed a conservative approach, relying primarily on tradi- tion and Science Department Program Assistant, Michael
tional animal classification schemes. Taxonomy has always been Souza, for his assistance during the project. The AZA is a pro-
a volatile field, with frequent arguments over the naming of or fessional membership association, representing 205 accred-
evolutionary relationships between various organisms. The ad- ited zoological parks and aquariums in North America. As
vent of DNA fingerprinting and other advanced biochemical Director/William Conway Chair, AZA Department of Con-
techniques has revolutionized the field and, not unexpectedly, servation and Science, I feel that I am a philosophical de-
has produced both advances and confusion. In producing these scendant of Dr. Grzimek, whose many works I have collected
volumes, we have consulted with specialists to obtain the most and read. The zoo and aquarium profession has come a long
up-to-date information possible, but knowing that new find- way since the 1970s, due, in part, to innovative thinkers such
ings may result in changes at any time. When scientific con- as Dr. Grzimek. I hope this latest revision of his work will
troversy over the classification of a particular animal or group continue his extraordinary legacy.
of animals existed, we did our best to point this out in the text.
Silver Spring, Maryland, 2001
Readers should note that it was impossible to include as
much detail on some animal groups as was provided on oth- Michael Hutchins
ers. For example, the marine and freshwater fish, with vast
numbers of orders, families, and species, did not receive as Series Editor

x Grzimek’s Animal Life Encyclopedia

•••••

How to use this book

Gzimek’s Animal Life Encyclopedia is an internationally As chapters narrow in focus, they become more tightly for-
prominent scientific reference compilation, first published in matted. General chapters have a loose structure, reminiscent
German in the late 1960s, under the editorship of zoologist of the first edition. While not strictly formatted, order chap-
Bernhard Grzimek (1909–1987). In a cooperative effort be- ters are carefully structured to cover basic information about
tween Gale and the American Zoo and Aquarium Association, member families. Monotypic orders, comprised of a single
the series is being completely revised and updated for the first family, utilize family chapter organization. Family chapters
time in over 30 years. Gale is expanding the series from 13 are most tightly structured, following a prescribed format of
to 17 volumes, commissioning new color images, and updat- standard rubrics that make information easy to find and un-
ing the information while also making the set easier to use. derstand. Family chapters typically include:
The order of revisions is:
Thumbnail introduction
Vol 8–11: Birds I–IV Common name
Vol 6: Amphibians Scientific name
Vol 7: Reptiles Class
Vol 4–5: Fishes I–II Order
Vol 12–16: Mammals I–V Suborder
Vol 1: Lower Metazoans and Lesser Deuterostomes Family
Vol 2: Protostomes Thumbnail description
Vol 3: Insects Size
Vol 17: Cumulative Index Number of genera, species
Habitat
Organized by order and family Conservation status

The overall structure of this reference work is based on Main essay
the classification of animals into naturally related groups, a Evolution and systematics
discipline known as taxonomy—the science through which Physical characteristics
various organisms are discovered, identified, described, Distribution
named, classified, and catalogued. Starting with the simplest Habitat
life forms, the protostomes, in Vol. 1, the series progresses Behavior
through the more complex animal classes, culminating with Feeding ecology and diet
the mammals in Vols. 12–16. Volume 17 is a stand-alone cu- Reproductive biology
mulative index. Conservation status
Significance to humans
Organization of chapters within each volume reinforces
the taxonomic hierarchy. Opening chapters introduce the Species accounts
class of animal, followed by chapters dedicated to order and Common name
family. Species accounts appear at the end of family chapters. Scientific name
To help the reader grasp the scientific arrangement, each type Subfamily
of chapter has a distinctive color and symbol: Taxonomy
Other common names
▲= Family Chapter (yellow background) Physical characteristics
Distribution
● = Order Chapter (blue background) Habitat
Behavior
●▲ = Monotypic Order Chapter (green background) Feeding ecology and diet
Reproductive biology
Grzimek’s Animal Life Encyclopedia
xi

How to use this book sible. For Conservation status, Grzimek’s follows the IUCN
Red List system, developed by its Species Survival Commis-
Conservation status sion. The Red List provides the world’s most comprehensive
Significance to humans inventory of the global conservation status of plants and an-
Resources imals. Using a set of criteria to evaluate extinction risk, the
Books IUCN recognizes the following categories: Extinct, Extinct
Periodicals in the Wild, Critically Endangered, Endangered, Vulnerable,
Organizations Conservation Dependent, Near Threatened, Least Concern,
Other and Data Deficient. For a complete explanation of each cat-
egory, visit the IUCN web page at http://www.iucn.org/
Color graphics enhance understanding themes/ssc/redlists/categor.htm

Grzimek’s features approximately 3,500 color photos, in- In addition to IUCN ratings, essays may contain other con-
cluding approximately 480 in four Birds volumes; 3,500 total servation information, such as a species’ inclusion on one of
color maps, including almost 1,500 in the four Birds volumes; three Convention on International Trade in Endangered
and approximately 5,500 total color illustrations, including Species (CITES) appendices. Adopted in 1975, CITES is a
1,385 in four Birds volumes. Each featured species of animal global treaty whose focus is the protection of plant and ani-
is accompanied by both a distribution map and an illustration. mal species from unregulated international trade.

All maps in Grzimek’s were created specifically for the pro- Grzimek’s provides the following standard information on
ject by XNR Productions. Distribution information was pro- avian lineage in Taxonomy rubric of each Species account:
vided by expert contributors and, if necessary, further [First described as] Muscicapa rufifrons [by] Latham, [in] 1801,
researched at the University of Michigan Zoological Museum [based on a specimen from] Sydney, New South Wales, Aus-
library. Maps are intended to show broad distribution, not tralia. The person’s name and date refer to earliest identifi-
definitive ranges, and are color coded to show resident, breed- cation of a species, although the species name may have
ing, and nonbreeding locations (where appropriate). changed since first identification. However, the organism de-
scribed is the same.
All the color illustrations in Grzimek’s were created specif-
ically for the project by Michigan Science Art. Expert con- Other common names in English, French, German, and
tributors recommended the species to be illustrated and Spanish are given when an accepted common name is avail-
provided feedback to the artists, who supplemented this in- able.
formation with authoritative references and animal skins from
University of Michgan Zoological Museum library. In addi- Appendices and index
tion to species illustrations, Grzimek’s features conceptual
drawings that illustrate characteristic traits and behaviors. For further reading directs readers to additional sources
of information about birds. Valuable contact information for
About the contributors Organizations is also included in an appendix. While the en-
cyclopedia minimizes scientific jargon, it also provides a Glos-
The essays were written by expert contributors, including sary at the back of the book to define unfamiliar terms. An
ornithologists, curators, professors, zookeepers, and other exhaustive Aves species list records all known species of birds,
reputable professionals. Grzimek’s subject advisors reviewed categorized according to Peters Checklist (1934–1986). And
the completed essays to insure that they are appropriate, ac- a full-color Geologic time scale helps readers understand pre-
curate, and up-to-date. historic time periods. Additionally, each of the four volumes
contains a full Subject index for the Birds subset.
Standards employed
Acknowledgements
In preparing these volumes, the editors adopted a conser-
vative approach to taxonomy, relying primarily on Peters Gale would like to thank several individuals for their im-
Checklist (1934–1986)—a traditional classification scheme. portant contributions to the series. Michael Souza, Program
Taxonomy has always been a volatile field, with frequent ar- Assistant, Department of Conservation and Science, Ameri-
guments over the naming of or evolutionary relationships be- can Zoo and Aquarium Association, provided valuable behind-
tween various organisms. The advent of DNA fingerprinting the-scenes research and reliable support at every juncture of
and other advanced biochemical techniques has revolution- the project. Also deserving of recognition are Christine Shep-
ized the field and, not unexpectedly, has produced both ad- pard, Curator of Ornithology at Bronx Zoo, and Barry Tay-
vances and confusion. In producing these volumes, Gale lor, professor at the University of Natal, in Pietermaritzburg,
consulted with noted taxonomist Professor Walter J. Bock as South Africa, who assisted subject advisors in reviewing man-
well as other specialists to obtain the most up-to-date infor- uscripts for accuracy and currency. And, last but not least,
mation possible. When scientific controversy over the classi- Janet Hinshaw, Bird Division Collection Manager at the Uni-
fication of a particular animal or group of animals existed, the versity of Michigan Museum of Zoology, who opened her col-
text makes this clear. lections to Grzimek’s artists and staff and also compiled the
“For Further Reading” bibliography at the back of the book.
Grzimek’s has been designed with ready reference in mind
and the editors have standardized information wherever fea- Grzimek’s Animal Life Encyclopedia

xii

•••••

Advisory boards

Series advisor Director, Marine Laboratory & Facilities
Humboldt State University
Michael Hutchins, PhD Arcata, California
Director of Conservation and William Conway Chair
American Zoo and Aquarium Association Volume 6: Amphibians
Silver Spring, Maryland William E. Duellman, PhD
Curator of Herpetology Emeritus
Subject advisors Natural History Museum and Biodiversity Research Center
University of Kansas
Volume 1: Lower Metazoans and Lesser Deuterostomes Lawrence, Kansas
Dennis Thoney, PhD
Director, Marine Laboratory & Facilities Volume 7: Reptiles
Humboldt State University James B. Murphy, PhD
Arcata, California Smithsonian Research Associate
Department of Herpetology
Volume 2: Protostomes National Zoological Park
Dennis Thoney, PhD Washington, DC
Director, Marine Laboratory & Facilities
Humboldt State University Volumes 8–11: Birds I–IV
Arcata, California Walter J. Bock, PhD
Permanent secretary, International Ornithological Congress
Sean F. Craig, PhD Professor of Evolutionary Biology
Assistant Professor, Department of Biological Sciences Department of Biological Sciences,
Humboldt State University Columbia University
Arcata, California New York, New York

Volume 3: Insects Jerome A. Jackson, PhD
Art Evans, PhD Program Director, Whitaker Center for Science,
Entomologist Mathematics, and Technology Education
Richmond, Virginia Florida Gulf Coast University
Ft. Myers, Florida
Rosser W. Garrison, PhD
Systematic Entomologist, Los Angeles County Volumes 12–16: Mammals I–V
Los Angeles, California Valerius Geist, PhD
Professor Emeritus of Environmental Science
Volumes 4–5: Fishes I–II University of Calgary
Paul Loiselle, PhD Calgary, Alberta
Curator, Freshwater Fishes Canada
New York Aquarium
Brooklyn, New York Devra Gail Kleiman, PhD
Smithsonian Research Associate
Dennis Thoney, PhD National Zoological Park
Washington, DC

Grzimek’s Animal Life Encyclopedia xiii

Advisory boards Charles Jones
Library Media Specialist
Library advisors Plymouth Salem High School
Plymouth, Michigan
James Bobick
Head, Science & Technology Department Ken Kister
Carnegie Library of Pittsburgh Reviewer/General Reference teacher
Pittsburgh, Pennsylvania Tampa, Florida

Linda L. Coates Richard Nagler
Associate Director of Libraries Reference Librarian
Zoological Society of San Diego Library Oakland Community College
San Diego, California Southfield Campus
Southfield, Michigan
Lloyd Davidson, PhD
Life Sciences bibliographer and head, Access Services Roland Person
Seeley G. Mudd Library for Science and Engineering Librarian, Science Division
Evanston, Illinois Morris Library
Southern Illinois University
Thane Johnson Carbondale, Illinois
Librarian
Oaklahoma City Zoo
Oaklahoma City, Oklahoma

xiv Grzimek’s Animal Life Encyclopedia

•••••

Contributing writers

Birds I–IV Donald F. Bruning, PhD Francis Hugh John Crome, BSc
Wildlife Conservation Society Consultant
Michael Abs, Dr. rer. nat. Bronx, New York Atheron, Queensland
Berlin, Germany Australia

George William Archibald, PhD Joanna Burger, PhD Timothy Michael Crowe, PhD
International Crane Foundation Rutgers University University of Cape Town
Baraboo, Wisconsin Piscataway, New Jersey Rondebosch, South Africa

Helen Baker, PhD Carles Carboneras H. Sydney Curtis, BSc
Joint Nature Conservation Committee SEO/BirdLife Queensland National Parks &
Peterborough, Cambridgeshire Barcelona, Spain Wildlife Service (Retired)
United Kingdom Brisbane, Queensland
John Patrick Carroll, PhD Australia
Cynthia Ann Berger, MS University of Georgia
Pennsylvania State University Athens, Georgia S. J. J. F. Davies, ScD
State College, Pennsylvania Curtin University of Technology
Robert Alexander Cheke, PhD Department of Environmental Biology
Matthew A. Bille, MSc Natural Resources Institute Perth, Western Australia
Colorado Springs, Colorado University of Greenwich Australia
Chatham, Kent
Walter E. Boles, PhD United Kingdom Gregory J. Davis, PhD
Australian Museum University of Wisconsin-Green Bay
Sydney, New South Wales Jay Robert Christie, MBA Green Bay, Wisconsin
Australia Racine Zoological Gardens
Racine, Wisconsin William E. Davis, Jr., PhD
Carlos Bosque, PhD Boston University
Universidad Simón Bolivar Charles T. Collins, PhD Boston, Massachusetts
Caracas, Venezuela California State University
Long Beach, California Stephen Debus, MSc
David Brewer, PhD University of New England
Research Associate Malcolm C. Coulter, PhD Armidale, New South Wales
Royal Ontario Museum IUCN Specialist Group on Storks, Australia
Toronto, Ontario Ibises and Spoonbills
Canada Chocorua, New Hampshire Michael Colin Double, PhD
Australian National University
Daniel M. Brooks, PhD Adrian Craig, PhD Canberra, A.C.T.
Houston Museum of Natural Science Rhodes University Australia
Houston, Texas Grahamstown, South Africa
Rachel Ehrenberg, MS
Grzimek’s Animal Life Encyclopedia University of Michigan
Ann Arbor, Michigan

xv

Contributing writers Frank Hawkins, PhD Jiro Kikkawa, DSc
Conservation International Professor Emeritus
Eladio M. Fernandez Antananarivo, Madagascar University of Queensland,
Santo Domingo Brisbane, Queensland
Dominican Republic David G. Hoccom, BSc Australia
Royal Society for the Protection of
Simon Ferrier, PhD Birds Margaret Field Kinnaird, PhD
New South Wales National Parks and Sandy, Bedfordshire Wildlife Conservation Society
Wildlife Service United Kingdom Bronx, New York
Armidale, New South Wales
Australia Peter Andrew Hosner Guy M. Kirwan, BA
Cornell University Ornithological Society of the Middle
Kevin F. Fitzgerald, BS Ithaca, New York East
South Windsor, Connecticut Sandy, Bedfordshire
Brian Douglas Hoyle PhD United Kingdom
Hugh Alastair Ford, PhD Bedford, Nova Scotia
University of New England Canada Melissa Knopper, MS
Armidale, New South Wales Denver Colorado
Australia Julian Hughes
Royal Society for the Protection of Niels K. Krabbe, PhD
Joseph M. Forshaw Birds University of Copenhagen
Australian Museum Sandy, Bedfordshire Copenhagen, Denmark
Sydney, New South Wales United Kingdom
Australia James A. Kushlan, PhD
Robert Arthur Hume, BA U.S. Geological Survey
Bill Freedman, PhD Royal Society for the Protection of Smithsonian Environmental Research
Department of Biology Birds Center
Dalhousie University Sandy, Bedfordshire Edgewater, Maryland
Halifax, Nova Scotia United Kingdom
Canada Norbert Lefranc, PhD
Gavin Raymond Hunt, PhD Ministère de l’Environnement,
Clifford B. Frith, PhD University of Auckland Direction Régionale
Honorary research fellow Auckland, New Zealand Metz, France
Queensland Museum
Brisbane, Australia Jerome A. Jackson, PhD P. D. Lewis, BS
Florida Gulf Coast University Jacksonville Zoological Gardens
Dawn W. Frith, PhD Ft. Myers, Florida Jacksonville, Florida
Honorary research fellow
Queensland Museum Bette J. S. Jackson, PhD Josef H. Lindholm III, BA
Brisbane, Australia Florida Gulf Coast University Cameron Park Zoo
Ft. Myers, Florida Waco, Texas
Peter Jeffery Garson, DPhil
University of Newcastle Darryl N. Jones, PhD Peter E. Lowther, PhD
Newcastle upon Tyne Griffith University Field Museum
United Kingdom Queensland, Australia Chicago, Illinois

Michael Gochfeld, PhD, MD Alan C. Kemp, PhD Gordon Lindsay Maclean, PhD, DSc
UMDNJ-Robert Wood Johnson Naturalists & Nomads Rosetta, South Africa
Medical School Pretoria, South Africa
Piscataway, New Jersey Steve Madge
Angela Kay Kepler, PhD Downderry, Torpoint
Michelle L. Hall, PhD Pan-Pacific Ecological Consulting Cornwall
Australian National University Maui, Hawaii United Kingdom
School of Botany and Zoology
Canberra, A.C.T. Grzimek’s Animal Life Encyclopedia
Australia

xvi

Contributing writers

Albrecht Manegold James David Rising, PhD Walter Sudhaus, PhD
Institut für Biologie/Zoologie University of Toronto Institut für Zoologie
Berlin, Germany Department of Zoology Berlin, Germany
Toronto, Ontario
Jeffrey S. Marks, PhD Canada J. Denis Summers-Smith, PhD
University of Montana Cleveland, North England
Missoula, Montana Christopher John Rutherford Robertson United Kingdom
Wellington, New Zealand
Juan Gabriel Martínez, PhD Barry Taylor, PhD
Universidad de Granada Peter Martin Sanzenbacher, MS University of Natal
Departamento de Biologia USGS Forest & Rangeland Ecosystem Pietermaritzburg, South Africa
Animal y Ecologia Science Center
Granada, Spain Corvallis, Oregon Markus Patricio Tellkamp, MS
University of Florida
Barbara Jean Maynard, PhD Matthew J. Sarver, BS Gainesville, Florida
Laporte, Colorado Ithaca, New York
Joseph Andrew Tobias, PhD
Cherie A. McCollough, MS Herbert K. Schifter, PhD BirdLife International
PhD candidate, University of Texas Naturhistorisches Museum Cambridge
Austin, Texas Vienna, Austria United Kingdom

Leslie Ann Mertz, PhD Richard Schodde PhD, CFAOU Susan L. Tomlinson, PhD
Fish Lake Biological Program Australian National Wildlife Texas Tech University
Wayne State University Collection, CSIRO Lubbock, Texas
Biological Station Canberra, A.C.T.
Lapeer, Michigan Australia Donald Arthur Turner, PhD
East African Natural History Society
Derek William Niemann, BA Karl-L. Schuchmann, PhD Nairobi, Kenya
Royal Society for the Protection of Alexander Koenig Zoological Research
Birds Institute and Zoological Museum Michael Phillip Wallace, PhD
Sandy, Bedfordshire Bonn, Germany Zoological Society of San Diego
United Kingdom San Diego, California
Tamara Schuyler, MA
Malcolm Ogilvie, PhD Santa Cruz, California John Warham, PhD, DSc
Glencairn, Bruichladdich University of Canterbury
Isle of Islay Nathaniel E. Seavy, MS Christchurch, New Zealand
United Kingdom Department of Zoology
University of Florida Tony Whitehead, BSc
Penny Olsen, PhD Gainesville, Florida Ipplepen, Devon
Australian National University United Kingdom
Canberra, A.C.T. Charles E. Siegel, MS
Australia Dallas Zoo Peter H. Wrege, PhD
Dallas, Texas Cornell University
Jemima Parry-Jones, MBE Ithaca, New York
National Birds of Prey Centre Julian Smith, MS
Newent, Gloucestershire Katonah, New York
United Kingdom
Joseph Allen Smith
Colin Pennycuick, PhD, FRS Baton Rouge, Louisiana
University of Bristol
Bristol, United Kingdom

Grzimek’s Animal Life Encyclopedia xvii

•••••

Contributing illustrators

Drawings by Michigan Gillian Harris, BA
Science Art Jonathan Higgins, BFA, MFA
Amanda Humphrey, BFA
Joseph E. Trumpey, Director, AB, MFA Jacqueline Mahannah, BFA, MFA
Science Illustration, School of Art and Design, University John Megahan, BA, BS, MS
of Michigan Michelle L. Meneghini, BFA, MFA
Bruce D. Worden, BFA
Wendy Baker, ADN, BFA
Brian Cressman, BFA, MFA Thanks are due to the University of Michigan, Museum
Emily S. Damstra, BFA, MFA of Zoology, which provided specimens that served as models
Maggie Dongvillo, BFA for the images.
Barbara Duperron, BFA, MFA
Dan Erickson, BA, MS Laura Exner
Patricia Ferrer, AB, BFA, MFA Andy Grosvold
Cory Johnson
Maps by XNR Productions Paula Robbins

Paul Exner, Chief cartographer
XNR Productions, Madison, WI

Tanya Buckingham
Jon Daugherity

xviii Grzimek’s Animal Life Encyclopedia

▲

Old World warblers

(Sylviidae)

Class Aves
Order Passeriformes
Suborder Passeri (Oscines)
Family Sylviidae

Thumbnail description
Very small to medium-sized, often dull-colored,
songbirds with thin, pointed bills

Size
3.1–9.8 in (8–25 cm); .1–2 oz (4–56 g)

Number of genera, species
60 genera; 350–391 species

Habitat Distribution
Highly varied. Largely arboreal, but many Subfamily Sylviinae: Palearctic region, Africa, Asia, Australasia, Oceania. Two
species inhabit wetlands, grasslands, thickets, genera, Regulus and Phylloscopus, reach Nearctic. Subfamily Polioptilinae: New
scrub, and riparian zones World, from South America to the northern United States.

Conservation status
Endangered: 10 species; Critically Endangered:
3 species; Vulnerable: 29 species; Near
Threatened: 13 species; Data Deficient: 9
species

Evolution and systematics are considered to be within the sylviid subfamily Sylviinae.
These are: (1) the kinglets (Regulus), regarded as a separate
The taxonomy of the Passeri, the suborder of oscine family by Sibley and Ahlquist and others; (2) the cisticolas and
passerines (the songbirds) that contains the Sylviidae (Old allies (Cisticola, Prinia, Apalis, Camaroptera), and other allied
World warblers), has long been debated. Much of the con- genera, a distinct group that Sibley and Ahlquist regard as a
troversy focuses on the delineation of the apparently closely separate family, and most authors regard as a subfamily, Cis-
related families Muscicapidae, Turdidae, Timaliidae, and ticolinae, of the Sylviidae; (3) the Sylvia warblers (Sylvia, Pari-
Sylviidae. Currently available molecular data suggest that soma), a group that, according to Sibley and Ahlquist, is more
widely used, traditional family classifications do not represent closely related to the timaliine babblers than to traditional
the evolutionary history of this large and complex group of sylviids; (4) the grassbirds and allies (Megalurus, Bowdleria,
birds. Sibley and Ahlquist place the Sylviidae within a large Cincloramphus, Megalurulus, Chaetornis, Gramnicola, Schoeni-
superfamily, the Sylvioidea. This superfamily also contains cola), and other allied genera, which comprise the subfamily
members of the following traditional families: Sittidae Megalurinae in the Sibley and Ahlquist system; (5) the re-
(nuthatches), Certhiidae (creepers), Troglodytidae (wrens), maining genera of the traditional Sylviidae, most of which ap-
Paridae (tits), Aegithalidae (long-tailed tits), Hirundinidae pear to fall within a clade represented by Sibley and Ahlquist’s
(swallows), Pycnonotidae (bulbuls), Zosteropidae (white- Acrocephalinae subfamily. Groups three, four and five above,
eyes), and Timaliidae (babblers). The molecular data do not plus the Garrulax laughing thrushes comprise Sibley and
support the long-held belief that the Sylviidae are closely re- Ahlquist’s more restricted Sylviidae, hereafter referred to as
lated to the Turdidae (thrushes) and the Muscicapidae (Old Sylviidae sensu strictu.
World flycatchers), for these two families fall into a separate
superfamily, the Muscicapoidea. The data do, however, sup- Little is known about the evolutionary history of passerines.
port the inclusion of some of the babblers (Timaliinae) and The prevailing opinion has been that passerines arose in the
laughing thrushes (genus Garrulax) in the Sylviidae family. Tertiary, specifically in the early Eocene, then underwent a
dramatic diversification during the Miocene. The oldest puta-
The classification used in the present work is based upon tive passerine fossils are from early Eocene Australia (ca. 54 mil-
the traditional definition of the Sylviidae. The genera may be lion years ago), lending support to a Southern Hemisphere
divided into six groups, based on molecular evidence of their origin, since passerine fossils do not appear in the Northern
taxonomic affiliation. The first of these are the gnatcatchers Hemisphere until the Oligocene. By the lower Miocene, passer-
and gnatwrens (Polioptila, Microbates, and Ramphocaenus), ine fossils greatly outnumber all other taxa in many Northern
which comprise the Sylviid subfamily Polioptilinae, but are Hemisphere sites. A recent molecular study suggests that the
placed by Sibley and Ahlquist within an expanded Certhiidae, passerine divergence may be much older and that passerines
including both creepers and wrens. The remaining five groups
1
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Family: Old World warblers 2 Vol. 11: Birds IV

1 3

4 56

Warbler nests: 1. Great reed warbler (Acrocephalus arundinaceus); 2. Common tailorbird (Orthotomus sutorius); 3. Blue-gray gnatcatcher (Po-
lioptila caerulea); 4. Fan-tailed cisticola (Cisticola juncidis); 5. Long-billed crombec (Sylvietta rufescens); 6. Tawny-flanked prinia (Prinia subflava).
(Illustration by Barbara Duperron)

evolved on Gondwana, the Cretaceous supercontinent in the flycatchers and thrushes); rictal bristles at the base of the bill
southern hemisphere. If this is the case, oscine passerines may (a widespread adaptation for insectivory); thin, pointed bill;
have diverged from suboscine passerines when Australia sepa- hatchlings naked or only partially downy; ten primaries;
rated from Gondwana, radiated on Australia, then dispersed scutellate tarsi; and lack of strong sexual dimorphism (Sylvia
throughout the Old World when Australia came in contact with is a notable exception). Most genera have twelve tail feathers,
Southeast Asia. but there are a few exceptions. Wing shape and length ranges
from short and rounded in sedentary species, to longer and
Within the Sylviidae sensu strictu, the first divergence was more pointed in long-distance migrants. Moult timing and
probably between Sibley & Ahlquist’s Acrocephaline sub- pattern are highly variable.
family (Acrocephalus, Hippolais, Phylloscopus, Seicercus, Sylvietta,
Sphenoeacus, etc.) and the lineage leading to the other three Sylviids are typically dull in color; often in shades of brown,
subfamilies. Next to diverge was Garrulax, the laughing- green, yellow, and gray. The family includes some of the tini-
thrushes, which diverged from a lineage leading to the Timali- est songbirds in the world, the kinglets, as well as some small
ine babbers and Sylvia and Parisoma warblers. The genus wren-like birds, many small warblers, and the medium-sized
Sylvia is thought to be at least 12-16 million years old. The marsh warblers and grassbirds. The smallest sylviids, the
Phylloscopus radiation is about as old as Sylvia, while the kinglets, weigh only a few grams. The largest, the marsh war-
Acrocephalus/Hippolais radiation is only about half as old. These blers Acrocephalus and the grassbirds Megalurus can weigh
estimates place much of the sylviid radiation during the close to 2 oz (60 g). A comprehensive summary of longevity
Miocene and Pliocene epochs of the mid-Tertiary, consistent data is unavailable, but many species live at least 8–12 years.
with the passerine radiation in the fossil record.
While most species do not have distinctive breeding
Physical characteristics plumages like those of the New World warblers (Parulidae),
a few sylviids show a marked contrast between breeding and
The traits that have been used to characterize Sylviidae nonbreeding plumages. A striking example is the red-winged
sensu latu are: unspotted young (as opposed to the Old World warbler (Heliolais erythroptera), the only member of a genus
apparently closely related to Prinia. While breeding red-
2
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Vol. 11: Birds IV Family: Old World warblers

A grasshopper warbler’s (Locustella naevia) wing-waving display. (Illustration by Barbara Duperron)

winged warblers of both sexes have dark gray upperparts and occurred between Northern (Palearctic) and Southern (Orien-
tail, they become predominantly tawny-brown in nonbreed- tal and African) Hemisphere faunas in the Old World, perhaps
ing plumage. A somewhat less striking example is the genus due to the East/West orientation of major barriers, including
Cisticola. Members of this genus undergo two moults each sea- the Sahara and Gobi deserts, the Himalayas, the Alps, the At-
son, and most have shorter tails and slightly different plumage las, Caucasus, and other mountain ranges, and historically, the
coloration during the breeding season. A few members of the Tethys sea, which separated Africa from what is now the
genus Prinia have similar distinct seasonal plumage variation. Palearctic. Conversely, in the Americas, where the major bar-
riers are oriented North to South, many Neotropical migrant
Distribution families originated in South America, then spread north to the
Nearctic. Voous (1977) is of the opinion that all Old World
The Old World warblers are extraordinarily widespread, sylviids arose in either the Indo-African region or the Sino-
occuring on every continent except South America and Antarc- Himalayan region. While these are areas where the largest ra-
tica. If the Polioptilinae are included, there are representatives diations have taken place, it is not clear where the common an-
of Sylviidae sensu latu in South America as well. Most of the cestor of the modern Sylviidae arose.
sylviid diversity occurs in the African and Oriental faunal re-
gions, with a less diverse, but widespread group of species in Habitat
the Palearctic. Nearctic and Australian species represent re-
cent invasions from Siberia and Southeast Asia, respectively. The Old World warblers occupy an astonishing variety of
habitats, from montane and riparian forests to arid scrublands
The centers of distribution for some major sylviid genera are to marshes and river floodplains to city parks and backyards.
as follows. Sino-Himalayan Region: Bradypterus, Cettia, Phyllo- Sylviids are found in all extremes of water availability with
scopus, Seicercus, Prinia; Southeast Asia: Megalurus and allies, Or- the exception of open water and harsh deserts. The family is
thotomus; Temperate Asia (Palearctic): Locustella, Acrocephalus, represented at a wide range of altitudes, from lowlands at or
Hippolais; Africa: Cisticola and allies; Mediterranean/Middle near sea level to montane forests and dwarf scrub as high as
Eastern (Palearctic) region: Sylvia. Relatively little exchange has several thousand meters. Considerable habitat adaptability is

Grzimek’s Animal Life Encyclopedia 3

Family: Old World warblers Vol. 11: Birds IV

Blue-gray gnatcatcher (Polioptila caerulea) chicks in their nest. (Photo tive and skulking. Most species inhabit dense vegetation, and
by R. & S. Day/VIREO. Reproduced by permission.) are most easily located and identified by voice. The carriage
of many of the Old World warblers is more or less horizon-
demonstrated by the varied habitats occupied by Acrocephalus tal. Flight ranges from very weak in some sedentary Afrotrop-
warblers that have colonized oceanic islands. ical species, to strong and sustained in long-distance migrants.
The gait is typically a hop, but some species run.
Habitat partitioning is widespread among sylviid warblers.
An example of foraging height segregation is found in the Vocalizations
Sylvietta crombecs. The long-billed crombec is restricted to The Old World warblers are one of the most vocally di-
undergrowth in areas of sympatry with red-faced and red-
capped crombecs. In other areas, the long-billed crombec is verse bird families in the world. Most species have well-
found at higher levels in the forest, demonstrating that com- developed songs, with parameters that vary greatly both
petition limits the ability of this species to fully exploit its po- within and between genera. Song ranges from the unmusical,
tential niche. Habitat partitioning also occurs in syntopic repetitive sounds of Prinia to the melodious warbles of Sylvia
species of Sylvia, but there is considerable overlap, leading to and Phylloscopus, to the amazingly complex, multi-element
interspecific territorial interactions. songs of some Acrocephalus species.

Exploitation of topological niches within a habitat has been Song is an important component of territorial defense; and
suggested as an important step in ecological and morpholog- the variety of habitats occupied by sylviids have probably con-
ical divergence of closely related species. Adam Richman and tributed to their vocal diversity. Species inhabiting relatively
Trevor Price have shown, in a 1992 study, that such a sce- open grasslands, marshes, and brushlands often engage in
nario has apparently occurred among a group of sympatric song-flight displays. Daily and seasonal singing patterns are
Phylloscopus warblers in the Himalayas. A related phenome- highly varied as well; Cettia cetti sings all day, for most of the
non is replacement, the presence of closely related species in year, sometimes at night. Night singing is common in Acro-
different habitat types, with little overlap. Replacement is es- cephalus, Bradypterus baboecala and Locustella.
sentially habitat partitioning on a larger scale. Many Afrotrop-
ical genera contain closely-related species that occupy similar Most Sylvia warblers appear to use subsong, typically a
ecological niches in dissimilar habitats. longer, lower, more complex and melodious version of the
territorial song, in courtship, but it is also given on the win-
Behavior tering grounds. A few sylviids incorporate mimicry into their
songs as well. The marsh warbler Acrocephalus palustris learns
Sylviids range from highly arboreal to almost entirely ter- its song components, from many dozens of other species it
restrial. They are generally very active but often quite secre- hears on both the breeding grounds in Europe, and the win-
tering grounds in Africa. Some of the Hippolais warblers (H.
4 icterina, H. languida, H. polyglotta) incorporate mimcry of other
species into phrases of their own songs. Mimicry also occurs
in Eremomela and Sylvia.

Duets are thought to be involved in pair-bond mainte-
nance, synchronization of sexual activity, or cooperative ter-
ritorial defense, and are generally associated with monogamous
species that maintain territories year-round. A Prinia subflava
female occasionally adds her own complementary rattle to the
notes of a singing male, creating a duet. The female Prinia
bairdii is known to duet during territory advertisement. A few
Cisticola species (C. hunteri, C. chubbi, C. nigriloris) also engage
in duetting. Apalis flavida duets, but not for territory defense.
Other genera with species that duet include: Bathmocercus,
Bradypterus, Drymocichla, Schistolais, and Spiloptila.

Migration and dispersal
The Sylviidae exhibit a wide range of movement tenden-

cies. There is even considerable variation within some species,
like the fan-tailed warbler (Cisticola juncidis), blue-gray gnat-
catcher (Polioptila caerulea), and Cetti’s warbler (Cettia cetti).
Each of these species includes some populations that are mi-
gratory, and others that are mostly sedentary. The true long-
distance migrants are typically Palearctic species that escape
the northern winter by travelling to sub-Saharan Africa or
Southern Asia. These long-distance migrants undertake their
movements at night.

Many Asian species are altitudinal migrants, moving be-
tween elevations, including the Tesias, several Phylloscopus and

Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World warblers

Seicercus species, and many Cettia species. Many African Garden warbler (Sylvia borin) chicks are fed in their nest. (Photo by
species undertake local and semi-local movements to moister Jany Sauvanet. Photo Researchers, Inc. Reproduced by permission.)
areas during the dry season. Cisticola juncidis and other species
of southern affinity tend to erupt northward after mild win- The typical foraging mode is perched gleaning (also
ters. Philopatry (tendency to remain near the home area) pre- known as standpicking) from the ground or from vegetation.
dominates in the great reed warbler, Acrocephalus arundinaceus, Sallying and hawking of aerial insects is also used by many
and is predicted by genetic diversity. species, including the flycatcher warblers (genus Seicercus).
Bradypterus sylvaticus forages on the ground by scratching up
Territorial and social interactions litter and humus with its feet, or by disturbing the humus
Almost all Old World warblers are territorial. Typically, with its wings and tail, in a motion reminiscent of dust-
bathing. Tesias, Macrosphenus kretschmeri, and other terres-
the male defends a territory with song, display, and some- trial warblers feed in a similar manner. Even some Phylloscopus
times chasing and fighting. The majority of antagonistic be- warblers, members of a genus composed mostly of arboreal
havior in migratory species occurs in the early spring during species, have adapted to ground-feeding. Kemp’s longbill
initial territory settlement and mate acquisition. In several Macrosphenus kempi climbs about on undergrowth, probing
monogamous species the female shares in territorial defense, its bill into dead wood.
sometimes by antagonistic behavior, and occasionally by song
or duetting with her mate. Among polygynous species, fe- Reproductive biology
males sometimes defend feeding and nesting areas within a
male’s territory against intrusion from neighboring females. Most Old World warblers are either socially monogamous
Many African species defend territories year-round. Winter- or polygynous. Monogamous warbler species may have a pair
ing and breeding ground site fidelity is often high in migra- bond that lasts for either a single nesting, a single season, or
tory and resident species. Family groups are common in many more frequently the pair may remain monogamous for several
genera after the breeding season has ended. years. Polygyny occurs when a single male maintains pair bonds
with two or more females, each with her own nest and young.
Many species are interspecifically territorial. Hippolais icte- Classical polyandry (sex-role reversal) does not occur in this
rina aggressively defends its territory and nest from many family. Serial monogamy is fairly common among the sylviids,
other species. Others, such as Cisticola anonymus, are only ag- with pair bonds lasting for only a single nesting. This is the
gressive toward other species near nest. Some sylviids, such case in Cisticola juncidis, a species in which most males mate se-
as the barred warbler Sylvia nisoria, preferentially associate quentially with three or four females (as many as eleven) per
with certain species of shrikes (Lanius) whose vigilance and season. Despite the implications of the term, many socially
anti-predator aggression result in higher reproductive success monogamous species engage in extra-pair copulations in order
for the barred warbler. to enhance their reproductive success. Polygyny has been in-
tensively studied in Acrocephalus reed warblers. Other genera
Feeding ecology and diet with polygynous species include Cettia, Cisticola, and Phyllosco-
pus. The duration of the pair bond is in part determined by the
The diets of all members of the family consist primarily of movements of a species.
insects, other arthropods, and especially spiders. Other prey
items include crustaceans, snails, and harvestmen (Opiliones). 5
Sap (golden-crowned kinglet) and nectar (several African and
Asian species, especially Prinia) are occasionally consumed.
Prinia hodgsonii and Orthotomus sutorius are known to carry
pollen attached to the feathers of the throat and forehead; it
and other nectar-feeding species may be important pollinators
in the tropics. Some of the larger reed-warblers (including
Acrocephalus arundinaceus, A. rufesecens, and A. stentoreus) occa-
sionally take small frogs and fish. Young sylviids are fed almost
exclusively arthropods, usually soft-bodied larvae and small in-
sects, but in some cases receive berries as well. Variation in
prey size and type is found among sympatric foraging guilds.
During the pre-migratory period of Palearctic Sylvia warblers,
individuals shift their diet from largely insects to largely berries
and fruits in order to accumulate fat for migration.

Foraging strategies among the Old World warblers are di-
verse. Some species forage singly or in pairs, while others for-
age in groups of several family members or other conspecifics.
Still others readily join mixed-species foraging parties, espe-
cially in the nonbreeding season. Cisticola nigriloris forages in
groups of 5–8 birds, apparently keeping in contact by group-
singing. Many African and nontropical species, join foraging
parties in the nonbreeding season.

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Family: Old World warblers Vol. 11: Birds IV

A black-browed reed-warbler (Acrocephalus bistrigiceps) builds its nest. pendulous, attached to living leaves, woven into surrounding
(Photo by A.J. Knystautas/VIREO. Reproduced by permission.) grasses, or built as a lining inside an abandoned nest of an-
other species. Nests of the Orthotomus tailorbirds are famous
Courtship for their sewing abilities. The female pierces green leaves with
Some species have elaborate courtship displays that may her bill, then draws them together with ‘threads’ of plant fiber,
making several stitches, and sometimes knotting the ends of
or may not be accompanied by vocal activity. During the the fibers. The resulting sac is lined with spider silk and plant
courtship dance of Camaroptera brachyura, the male repeat- fibers. Similar nests are made by Artisornis, Calamonastes, Ca-
edly jumps 12–16 in (30–40 cm) into the air from his perch, maroptera, and Schistolais. The nests of most sylviids are built
while whirring his wings and giving a precisely timed call each by both sexes, but in some species, especially polygynous ones,
second, corresponding with the jumps. Some Cisticola species the female builds alone.
use similar courtship dances, while others attract females and
defend their territories with remarkable aerial song-displays. Care of young
Many species, including the wood warbler (Phylloscopus sibila- Sylviids generally lay between one and 12 eggs per clutch.
trix) and the melodious warbler (Hippolais polyglotta), use a spe-
cial flight style, often called flutter-flight or butterfly flight The young are usually incubated and brooded by the female,
when pursuing a female. A characteristic of Sylvia courtship but in some species including Abroscopus, Seicercus, and Sylvia,
is the building of several cock nests (partial nests decorated the male shares in the task. Feeding of the young is usually
with spider silk, flower petals, etc.) by the male. The female done by both parents in socially monogamous species, but
apparently uses these partial nests to assess male quality. there is much interspecific and intraspecific variation in the
Courtship displays are diverse, and utilize full song, subsong, degree of paternal involvement. In socially polygynous
calls, posture and flight displays, nest building, and presenta- species, males tend to take a smaller role in feeding the
tion of objects. young. Aquatic warbler (Acrocephalus paludicola) males take no
Nests role in incubation and rearing of their young. The altricial
young, which are usually naked but may be downy, remain
The Old World warblers build some of the most interest- in the nest for 10–21 days. In many species, the fledglings
ing nests in the avian world. They range from simple, open are dependent on their parents for an additional 1–4 weeks
cups to intricately woven domed and rounded structures, to after leaving the nest. Broods of fledglings often roost in
highly camouflaged balls of moss and lichens. Nests may be groups to conserve warmth and receive food from the par-
located on the ground, in low vegetation, or in shrubs and ents. Most northern species are usually single-brooded, but
trees as high as 80 ft (26 m) or more. Support may be pro- occasionally raise a second brood if conditions permit. Many
vided by grass and sedge clumps, tree branches, or other veg- tropical and sub-tropical species, on the other hand, are
etation. Alternatively, the nest may be hanging or nearly double-brooded.

6 Some sylviid taxa engage in cooperative breeding. Helpers
have been reported in Acrocephalus vaughani, Prinia bairdii, Cis-
ticola anonymous, C. nigriloris, and most of the Eremomelas. It
is not clear, however, whether the helpers in these species are
related to the breeding pair, nor is it known whether mating
of helpers with breeders occurs. The only known case of true
cooperative breeding (a monogamous pair helped by non-
breeding kin) among the Sylviidae is the Seychelles warbler
(Acrocephalus sechellensis). Most of the helpers are females, since
males tend to disperse and attempt to set up their own terri-
tories. In high quality territories, the presence 1–2 helpers in-
creases the reproductive success of the resident pair, while the
presence of 3 or more helpers creates too much competition
and decreases reproductive success. Breeding pairs have been
shown to facultatively manipulate the sex ratio of their eggs
to maximize their future reproductive success.

Conservation status

Several species of Sylviid face serious threats. Many of
these species are greatly range-restricted, with small popula-
tions inhabiting a single island or tiny patch of isolated habi-
tat. Nearly one third of these threatened species are found in
tropical Africa. About as many are endemic to small oceanic
islands. The genera with the largest number of threatened or
Near Threatened species are: Acrocephalus (15), Apalis (7),
Bradypterus (6), and Cisticola (5).

Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World warblers

The reed-warblers (Acrocephalus), are among the most An arctic warbler (Phylloscopus borealis) sings while perched on a
widespread of the Sylviid genera, but their colonization of is- branch. (Photo by Doug Wechsler/VIREO. Reproduced by permission.)
land habitats makes them vulnerable to anthropogenic and
climatic threats. Acrocephaline species that have large ranges This species inhabits lowland coastal forests (to 500m) of
are presumably able to survive loss of habitat. However, is- Brazil, Argentina, and Paraguay, primarily in the Paraná basin.
land populations, and those species with small or already frag- Its habitat has been severely reduced and deforestation con-
mented ranges are susceptible to habitat destruction and tinues to pose a threat to its remaining small habitat.
introduction of nonindigenous species. The aquatic warbler
has probably suffered dramatic population declines due to de- Significance to humans
struction of its riverine marshland habitat in Eastern Europe
and Western Siberia. IUCN lists it as Vulnerable. Their small size, inconspicuous coloration, and preference
for inhabiting vegetation make the Old World warblers un-
The Seychelles warbler is an island species whose popula- likely candidates for human attention. However, the songs of
tion had drooped precipitously to a mere 26 birds occupying many species are complex and well-developed. Since many
the small Cousin Island in the Seychelles. Intensive manage- species inhabit Europe and the Mediterranean, these songs have
ment of the island resulted in a dramatic increase over the been familiar to Western civilization for thousands of years.
past thirty years, with the island reaching apparent carrying
capacity in the early 1980s. Recent translocation efforts have Sylviids are largely insectivorous, and are thus invaluable
expanded the range of the Seychelles Warbler to include two agents of pest control, not only in agricultural and suburban
more islands. areas, but in evergreen and hardwood forests. These small
songbirds are part of a natural balance that man repeatedly
The millerbird (Acrocephalus familiaris) is the only Sylviid threatens with pesticide use, introduction of alien species, and
to colonize the Hawaiian Archipelago. Historically, two pop- planting and maintenance of monocultures in farming and
ulations, which were usually regarded as separate species or timber production. Along with spiders and other natural en-
subspecies, inhabited the islands, one on Laysan and one on emies, sylviid warblers are part of an already extant system of
Nihoa. The Laysan form became extinct in the early twenti- insect control awaiting utilization by man. In many tropical
eth century, after much of the native vegetation was destroyed areas, warblers are major consumers of mosquitos, helping to
by rabbits. The Nihoan form persists on the steep, rocky limit the number of vectors for insect-borne disease.
slopes of this tiny volcanic island. The Nihoan millerbird is
listed as a Federally Endangered species in the United States.
A 1996 USFWS population estimate was 155 individuals.

Many of the Endangered African species are found in
threatened East African forests. A bird that may once have in-
habited natural forest clearings, Artisornis moreaui, is limited
in its dispersal ability by very weak flight. Forest clearing in
its very small range may lead to the extinction of this highly
local and poorly known species. Six species of Apalis, most
found only in the highland forests of one or a few mountain
groups of East Africa from Kenya to Mozambique, face seri-
ous conservation risks, as do rare species of Bathmocercus,
Chloropeta, Cisticola and Eremomela.

Several threatened species in the genera Acrocephalus,
Bradypterus, Cettia, Megalurulus, Megalurus, Locustella, Phyllo-
scopus, and Orthotomus occur on islands in Southeast Asia and
Australasia.

Among the kinglets, there has been only a single subspecies
faced with serious conservation threats. Regulus calendula ob-
scura was endemic to a highly restricted area of cypress groves
on the island of Guadalupe. The species was last seen in the
1970s and is considered Extinct.

Among the Polioptilinae, only one species, the creamy-
bellied gnatcatcher, Polioptila lactea, is is Near Threatened.

Grzimek’s Animal Life Encyclopedia 7

1
2
3

4
6

5

7 9
8

10 11

1. Golden-spectacled warbler (Seicercus burkii); 2. Whitethroat (Sylvia communis); 3. Golden-crowned kinglet (Regulus satrapa); 4. Tawny-flanked
prinia (Prinia subflava); 5. Long-billed crombec (Sylvietta rufescens); 6. Blackcap (Sylvia atricapilla); 7. Slaty-bellied tesia (Tesia olivea); 8. Arctic
warbler (Phylloscopus borealis); 9. Long-billed gnatwren (Ramphocaenus melanurus); 10. Blue-gray gnatcatcher (Polioptila caerulea); 11. Chiffchaff
(Phylloscopus collybita). (Illustration by Barbara Duperron)

8 Grzimek’s Animal Life Encyclopedia

12 3

4 6
5
7
9 8

10

11 12

1. Zitting cisticola (Cisticola juncidis); 2. Cetti’s warbler (Cettia cetti); 3. Common tailorbird (Orthotomus sutorius); 4. Yellow-breasted apalis (Apalis
flavida); 5. Yellow-bellied eremomela (Eremomela icteropygialis); 6. Icterine warbler (Hippolais icterina); 7. Marsh warbler (Acrocephalus palus-
tris); 8. Little grassbird (Megalurus gramineus); 9. Great reed warbler (Acrocephalus arundinaceus); 10. Little rush-warbler (Bradypterus baboe-
cala); 11. Golden-headed cisticola (Cisticola exilis); 12. Grasshopper warbler (Locustella naevia). (Illustration by Barbara Duperron)

Grzimek’s Animal Life Encyclopedia 9

Family: Old World warblers Vol. 11: Birds IV

Species accounts

Long-billed gnatwren BEHAVIOR

Ramphocaenus melanurus Very active. Solitary or in pairs. Song is a clear musical trill.
Male tail fanning and lateral tail movements probably function
SUBFAMILY as territorial or mate-attraction displays.

Polioptilinae FEEDING ECOLOGY AND DIET

TAXONOMY Forages for insects in low undergrowth, sometimes joins mixed
foraging flocks.
Ramphocaenus melanurus Vieillot, 1819.
REPRODUCTIVE BIOLOGY
OTHER COMMON NAMES
Nest is loose, deep cup of leaves, grasses and other plant materi-
French: Microbate à long bec; German: Schwarzschwanz- als located near the ground. Both parents incubate the two eggs
Degenschnäbler; Spanish: Chirito Picón. (17 days), and feed the young. Fledging occurs after 12-15 days.

PHYSICAL CHARACTERISTICS CONSERVATION STATUS

4.75 in (12 cm); .3–.4 oz (8.5–10 g). A small, distinctive bird Not threatened.
with a long, cocked tail and long, pale slender bill. Tawny
brown above, whitish below, with buffy wash on face, sides of SIGNIFICANCE TO HUMANS
breast and flanks. Tail dusky black, with white-tipped outer
feathers. None known. ◆

DISTRIBUTION Blue-gray gnatcatcher

Neotropical. Lowlands (to 4,900 ft [1500 m]) from Yucatan Polioptila caerulea
south through Central America and northern South America to
central Brazil, also east coast of Brazil, west of Andes to coastal SUBFAMILY
Ecuador.
Polioptilinae
HABITAT
TAXONOMY
Undergrowth and thickets in deciduous forest and forest edges,
and humid forest interior; vine tangle. Polioptila caerulea Linnaeus, 1766. Seven to nine subspecies rec-
ognized.

Ramphocaenus melanurus Polioptila caerulea

Resident Resident Breeding Nonbreeding

10 Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World warblers

OTHER COMMON NAMES Regulus satrapa

English: Cat-bird; French: Gobe-moucherons Gris-bleu; Ger- Resident Breeding Nonbreeding
man: Blaumükenfänger; Spanish: Perlita Grisilla.
crown. Male has orange central crown feathers that are usually
PHYSICAL CHARACTERISTICS concealed.

4–4.5 in (10–11.5 cm); 0.18–0.25 oz (5–7 g). Tiny grayish bird DISTRIBUTION
with a distinct white eye ring, and a long tail, often cocked.
Upperparts blue-gray, underparts whitish, outer tail feathers Breeds from Nearctic boreal zone south through New England
white, inner tail feathers black. Breeding male has black ‘fore- and Appalachians in eastern United States, Rockies, Cascades,
head’ stripe extending from base of bill to just above and be- Sierra Nevada, Chiricahuas in West. Resident populations in
hind the eye. mountains of Mexico, Guatemala. Winters south to Northeast-
ern Mexico.
DISTRIBUTION
HABITAT
Nearctic. Breeds throughout much of United States and Mex-
ico south to Belize. Winters from extreme southern United Dense conifers above 6,560 ft (2,000 m) in Mexico. Sometimes
States to Baja, Honduras, and Cuba. in deciduous forest in winter.

HABITAT BEHAVIOR

Swampy deciduous or pine woods and riparian lowlands in Very active, often hangs upside down. Voice includes a high
eastern United States. Arid scrub, pinyon-juniper, and open thin call note and song, given by both sexes, consisting of a se-
woodland in western United States. Scrub, wood edges, thorn ries of ascending notes, sometimes followed by a descending
forest, and clearings on wintering grounds. warble. Male defends territory with song, and song-displays
(crown raised, tail and wing flicking). Often joins mixed-species
BEHAVIOR flocks in fall and winter.

Usually solitary or in pairs. Call is a thin buzzy whine, the FEEDING ECOLOGY AND DIET
male song, a rather soft series of such notes, interspersed with
chips and whistles. In migratory populations, males arrives first Mostly small insects, spiders, and arthropod eggs; occasionally
and sing to defend their territories. Some populations in Mex- sap, rarely fruit. Occasionally hover-gleans.
ico and Bahamas are permanent residents, but most are migra-
tory between April and September. REPRODUCTIVE BIOLOGY

FEEDING ECOLOGY AND DIET Serially monogamous. Pair bond is maintained through breed-
ing season. Nest, built by both sexes, is a deep hanging cup of
Will take a wide variety of insects, spiders. Usually gleans moss, lichens, bark, spider webs and other plant material, at-
while perched, but also hover-gleans and hawks for insects. tached to hanging twigs near trunk, placed high in conifer.
Eight to nine (sometimes 5-11) eggs. Incubation by female
REPRODUCTIVE BIOLOGY only (14-15 days). Nestlings are fed by both parents, and leave
the nest after 14-19 days.
Monogamous. During courtship, the male leads the female to
potential nest sites. The nest, built by both sexes, is a neat cup CONSERVATION STATUS
of plant fibers often camouflaged and placed high on a branch
or fork of a tree or shrub. Four or five eggs are incubated by Not threatened.
both parents for 11–15 days. Female broods the nestlings, but
later both sexes feed young. Fledging occurs after 10–15 days. 11

CONSERVATION STATUS

Not threatened.

SIGNIFICANCE TO HUMANS

None known. ◆

Golden-crowned kinglet

Regulus satrapa

SUBFAMILY

Sylviinae

TAXONOMY

Regulus satrapa Lichtenstein, MHK, 1823.

OTHER COMMON NAMES

French: Roitelet à couronne dorée; German: Satrap; Spanish:
Reyezuelo Corona Dorada.

PHYSICAL CHARACTERISTICS

3–4 in (8–11 cm); 0.1–0.3 oz (4–7.5 g). Among smallest of all
songbirds; olive-green with two whitish wing bars, and a white
eye line surmounted by black lateral crown stripes and a yellow

Grzimek’s Animal Life Encyclopedia

Family: Old World warblers Vol. 11: Birds IV

SIGNIFICANCE TO HUMANS REPRODUCTIVE BIOLOGY

None known. ◆ Serially monogamous with most males mating with 1–11 fe-
males over the course of a season. Occasionally simultaneously
Zitting cisticola polygynous. Pair bond lasts for a single nesting. During
courtship, male builds several partially complete nests near the
Cisticola juncidis ground, and attracts female with song-flight. Nest is pear-
shaped bag made by sewing and weaving grasses together with
SUBFAMILY spider web. Two to six eggs incubated by female for 13 days;
young leave nest in 11–15 days. Female feeds young 10–20
Sylviinae days after fledging.

TAXONOMY CONSERVATION STATUS

Sylvia juncidis Rafinesque, 1810. Not threatened.

OTHER COMMON NAMES SIGNIFICANCE TO HUMANS

English: Fantailed warbler, streaked cisticola; French: Cisticole None known. ◆
des joncs; German: Cistensänger; Spanish: Buitrón Común.
Golden-headed cisticola
PHYSICAL CHARACTERISTICS
Cisticola exilis
3.9–4.7 in (10–12 cm); 0.3–0.4 oz (8–12 g). Small warbler with
warm brown upperparts strongly streaked with black, rufous SUBFAMILY
rump and flanks, short, rounded wings, and short, graduated tail,
spotted black and white underneath. Bill short, thin, and slightly Sylviinae
decurved.
TAXONOMY
DISTRIBUTION
Cisticola exilis Vigors and Horsfield, 1827.
Widespread. Southern Europe (Iberian Peninsula, Mediter-
ranean rim), sub-Saharan Africa, Indian subcontinent, South- OTHER COMMON NAMES
east Asia, Australasia.
English: Bright-headed cisticola, yellow-headed cisticola, tailor-
HABITAT bird, corn bird; French: Cisticole à couronne dorée; German:
Goldkopg-Cistensänger; Spanish: Buitrón de Capa Dorada.
Open tall-grass habitat and grassy wetlands, agricultural lands,
primarily in lowlands. PHYSICAL CHARACTERISTICS

BEHAVIOR 3.9–4.3 in (10–11 cm). Small warbler, warm brown back, ru-
fous nape, crown, breast and flanks, whitish throat and belly.
Mostly sedentary, but marked post-breeding dispersal of both Strong black streaking on back. Wings and tail short. Bill
adults and juveniles in many populations. Also migratory in short, thin, and slightly decurved.
Western Mediterranean. Male song is a quick, sharp single
note given consistently every 0.5 to 1 seconds. Males are ag-
gressively territorial, especially in vicinity of nest.

FEEDING ECOLOGY AND DIET

Forages mostly on the ground. Takes insects and insect larvae,
particularly Lepidoptera, grasshoppers, spiders, and beetles.

Cisticola juncidis Cisticola exilis

Resident Breeding Resident

12 Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World warblers

DISTRIBUTION OTHER COMMON NAMES

The only cisticola absent from Africa. Southern Asia, from In- English: Plain prinia; French: Prinia modeste; German:
dia and China south through Phillipines, Malay Archipelago, Braunkopfprinie; Spanish: Prinia de Dorsos Castaños.
and New Guinea, to Australia.
PHYSICAL CHARACTERISTICS
HABITAT
5.1–5.5 in (13–14 cm); 0.3–0.4 oz (7–11.5 g). Mmedium to
Tall, coarse, open grassland. Often in wet areas. large warbler with short rounded wings and a fairly long, grad-
uated tail. Upperparts uniformly rufous-brown to gray-brown,
BEHAVIOR underparts whitish. Short whitish eye line. Legs long, bill rela-
tively short, slightly decurved.
Usually solitary or in pairs. Scurries through dense grasses in a
mouse-like manner. May form loose flocks in nonbreeding sea- DISTRIBUTION
son. Male song, given during flight-display, is a nasal wheeze,
followed by an explosive, liquid plook note. Sub-Saharan Africa, southern Asia, from Indian subcontinent
to China, Taiwan, most of mainland Southeast Asia.
FEEDING ECOLOGY AND DIET
HABITAT
Insectivorous. Forages on or near ground.
Lowlands; grassland and savanna, wetlands, mangroves, scrub,
REPRODUCTIVE BIOLOGY forest clearings and edges.

Polygynous. Males with shorter tails have increased reproduc- BEHAVIOR
tive success. Male flight-display consists of characteristic circu-
lar flight, singing and a high-speed vertical plummet. Nest is a Active, vocal and conspicuous. Often in pairs or family groups.
small, rounded bag of grasses, plant down, and spider silk, at- Males territorial, defending year-round by singing. Apparently
tached to grasses or other vegetation. The female builds the sedentary, but movement may occur in parts of range during
nest, sometimes with help from the male, but incubates the dry season. Duetting occasionally occurs.
3–4 eggs herself.
FEEDING ECOLOGY AND DIET
CONSERVATION STATUS
Forages in vegetation, gleaning insects and sometimes taking
Not threatened. nectar.

SIGNIFICANCE TO HUMANS REPRODUCTIVE BIOLOGY

None known. ◆ Nest is an oval of woven grasses with a side-top entrance. Two
to five eggs are incubated for 13–14 days. Young, fed by both
Tawny-flanked prinia parents, fledge after 13–17 days, but remain with parents 2–3
weeks.
Prinia subflava
CONSERVATION STATUS
SUBFAMILY
Not threatened.
Sylviinae
SIGNIFICANCE TO HUMANS
TAXONOMY
None known. ◆
Motacilla subflava Gmelin, 1789. Subspecies inornata Sykes,
1832. Yellow-breasted apalis

Prinia subflava Apalis flavida

Resident SUBFAMILY

Grzimek’s Animal Life Encyclopedia Sylviinae

TAXONOMY

Apalis flavida Strickland, 1852.

OTHER COMMON NAMES

English: Brown-tailed apalis; French: Apalis à gorge jaune;
German: Gelbbrust-Feinsänger; Spanish: Apalis de Pecho
Amarillo.

PHYSICAL CHARACTERISTICS

4.7–5.1 in (12–13 cm); 0.3 oz (7–9.5 g). Small warbler, olive-
green above, whitish throat and belly separated by broad yel-
low breast band, with a central black marking in some
populations. Tail long and graduated, wings short and
rounded. Gray on crown and sides of face.

DISTRIBUTION

Endemic to sub-Saharan Africa.

HABITAT

Savanna, forest edge, riparian forest and thickets.

13

Family: Old World warblers Vol. 11: Birds IV

Apalis flavida Tesia olivea

Resident Resident

BEHAVIOR PHYSICAL CHARACTERISTICS

Often in pairs or family groups. Male song individually distinc- 3.5–4 in (9–10 cm); 0.2–0.3 oz (6–9 g). Characterized by an ex-
tive. Pairs sing duets. Maintain year-round feeding territories. ceptionally short tail with 10 rectrices, a large head, rounded
wings, long legs, and a more or less upright posture.
FEEDING ECOLOGY AND DIET
DISTRIBUTION
Forages in foliage of trees and thickets. Takes a variety of in-
sects, insect larvae, spiders, and also fruit and nectar. Some- From south Central Asia through lower Himalayas and Nepal
times joins mixed-species foraging parties. to south China, south to northwest Thailand, north Laos, and
North Vietnam.
REPRODUCTIVE BIOLOGY
HABITAT
Nest is a hanging, domed bag with side-top entrance, con-
structed largely of lichen bound with spider silk, 3–6 ft (1–2 m) Damp areas and stream courses in evergreen forests; usually
above ground. Two to three eggs incubated 12–14 days by the 3,280–8,860 ft (1,000–2,700 m).
female; the young, fed by both sexes, fledge after 15–16 days.
BEHAVIOR
CONSERVATION STATUS
Keeps to dense cover near the ground. It is skulking, but read-
Not threatened. ily approaches to investigate and scold an intruder. Apparently
non-migratory, but descends to lower elevations in winter.
SIGNIFICANCE TO HUMANS Voice a ventriloquial tchirik-tchirik.

None known. ◆ FEEDING ECOLOGY AND DIET

Slaty-bellied tesia Forages in litter on the forest floor for insects and spiders.

Tesia olivea REPRODUCTIVE BIOLOGY

SUBFAMILY Little known. Pairs frequently observed.

Sylviinae CONSERVATION STATUS

TAXONOMY Not threatened.

Tesia olivea McClelland, 1840. SIGNIFICANCE TO HUMANS

OTHER COMMON NAMES None known. ◆

English: Slaty-bellied ground warbler; French: Té à ventre ar- Cetti’s warbler
doise; German: Goldscheiteltesia; Spanish: Trinador de Vien-
tre Pizarro. Cettia cetti

14 SUBFAMILY

Sylviinae

TAXONOMY

Cettia cetti Temminck, 1820.

Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World warblers

SIGNIFICANCE TO HUMANS

None known. ◆

Cettia cetti Breeding Nonbreeding Little rush warbler

Resident Bradypterus baboecala

OTHER COMMON NAMES SUBFAMILY

French: Bouscarle de Cetti; German: Seidensänger; Spanish: Sylviinae
Ruiseñor Bastardo de Cetti.
TAXONOMY
PHYSICAL CHARACTERISTICS
Bradypterus baboecala Vieillot, 1817.
5.3–5.7 in (13.5–14.5 cm); 0.4–0.6 oz (10–18g) (males), 0.3–0.6
oz (8–16g) (females). Medium-sized, plump warbler with long, OTHER COMMON NAMES
graduated, rounded tail with 10 rectrices; short, rounded
wings; and a delicate, dark bill. Strongly sexually dimorphic, as English: African bush-warbler, African sedge warbler; French:
measured by wing length and mass. Upperparts dull chestnut Bouscarle des marais; German: Sumpfbuschsänger; Spanish:
to rufous, throat white, breast grayish, and belly and flanks Ruiseñor Africano.
buffy. Characteristic thin, whitish supercilium and eye ring.
PHYSICAL CHARACTERISTICS
DISTRIBUTION
5.9–7.5 in (15–19 cm); 0.4–0.6 oz (11–17 g). Medium-sized
Distribution across Eurasia. In summer, north into Caucasus warbler with dark brown upperparts, buffy flanks and breast,
region of Russia and Uzbekistan, Kyrgyzstan, Kazakstan. Win- whitish underparts, whitish throat streaked or spotted with
ters along the Indus River in Pakistan. Eastern race (albiventris) dark brown, and pale eye line. Tail wide and rounded.
migrates to south to Pakistan and Afghanistan.
DISTRIBUTION
HABITAT
Sub-Saharan Africa. Widespread, but local in the northern
Lowlands, usually near water, in dense thickets and reedbeds. tropics; common throughout much of the south.

BEHAVIOR HABITAT

Skulking. Male territorial song is an explosive series of clear Reedbeds and grasses, near marshes, lagoons, sewage ponds,
tones. A softer variant is used during courtship. Male song pat- watercourses.
tern is unique allowing individual recognition. Males aggres-
sively territorial, defending with song, wing-waving display and BEHAVIOR
fighting. In sedentary populations, territories defended all year.
Found singly or in pairs. Reluctant to fly. Song an accelerating
FEEDING ECOLOGY AND DIET series of chips, with the tempo of a “bouncing ball.” Males de-
fend territory with song and song-flight. Generally sedentary.
Forages on or near the ground for insects (especially aquatic
invertebrates), spiders, harvestmen, snails, earthworms, and Bradypterus baboecala
some seeds.
Resident
REPRODUCTIVE BIOLOGY
15
Often serially polygynous. Male may mate with the same 1–4 fe-
males for several successive years. Nest is loose cup of stems and
leaves, placed low among tangled vegetation. The nest of each
female is placed in her ‘range’ within the male territory. The
4–5 eggs are incubated by the female. Fledging occurs at 14–16
days, young remain dependent for additional 15 or more days.

CONSERVATION STATUS

Not threatened.

Grzimek’s Animal Life Encyclopedia

Family: Old World warblers Vol. 11: Birds IV

FEEDING ECOLOGY AND DIET BEHAVIOR

Forages for insects. Also known to take ant eggs. Very secretive; keeps to dense cover. Gait is a run; flight short
and low. Song a high, far-carrying trill, common at night. Fe-
REPRODUCTIVE BIOLOGY male sings during courtship.

Monogamous. Nest is a tight, bulky cup of grasses and reeds, FEEDING ECOLOGY AND DIET
situated in a tussock and supported by sedge or rush leaves
over water. The 2–3 eggs are incubated 12–14 days. Fledging Forages in low vegetation and on ground for insects.
period is 12–13 days, both sexes feed young.
REPRODUCTIVE BIOLOGY
CONSERVATION STATUS
Female builds cup nest of grass and plant material on or near
Not threatened. ground in thick vegetation. The 5–6 eggs are incubated 12–15
days by both parents. Young leave nest after 10–15 days.
SIGNIFICANCE TO HUMANS
CONSERVATION STATUS
None known. ◆
Not threatened.
Grasshopper warbler
SIGNIFICANCE TO HUMANS
Locustella naevia
None known. ◆
SUBFAMILY
Marsh warbler
Sylviinae
Acrocephalus palustris
TAXONOMY
SUBFAMILY
Locustella naevia Boddaert, 1783.
Sylviinae
OTHER COMMON NAMES
TAXONOMY
French: Locustelle tachetée; German: Feldschwirl; Spanish:
Buscarla Pintoja. Acrocephalus palustris Bechstein, 1798.

PHYSICAL CHARACTERISTICS OTHER COMMON NAMES

4.7–5.1 in (12–13 cm); 0.3–0.6 oz (9.5–18 g). Small grass war- French: Rousserolle verderolle; German: Sumpfrohrsänger;
bler with olive-brown upperparts, streaked with black, under- Spanish: Carricero Poliglota.
parts creamy white, flanks and breast buffy, streaked brown.
Short, rounded wings and short graduated tail. PHYSICAL CHARACTERISTICS

DISTRIBUTION 5.1 in (13 cm); 0.4–0.7 oz (10–20 g). Heavy, medium-sized
warbler with uniform olive-brown upperparts, creamy under-
Breeds from southern Scandinavia, British Isles and France parts, buffy wash on sides of breast and flanks, white throat,
west throughout central Europe and Siberia to Mongolia, and light line from eye to base of bill.
Northern China. Winters in Africa and Indian subcontinent.
DISTRIBUTION
HABITAT
Breeds throughout Central and Western Europe, excluding
Grasslands, low scrub, bogs, fens and marshes, with some Iberia and British Isles to southern Scandinavia and western
shrubs and trees. Russia. Winters in Southeast Africa.

Locustella naevia Acrocephalus palustris

Breeding Nonbreeding Breeding Nonbreeding

16 Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World warblers

HABITAT PHYSICAL CHARACTERISTICS

Breeds in tall herbaceous vegetation and woody cover, often in 7.5–7.9 in (19–20 cm); 0.7–1.8 oz (21–51 g). Large, heavy war-
moist areas, but also on dry slopes, and in parks and open forest. bler with strong bill and legs, longish pointed wings. Uniform
brown above, whitish below, with buffy wash on flanks, rufous
BEHAVIOR rump, light eyeline.

Solitary and territorial in breeding season, some males polyter- DISTRIBUTION
ritorial. Posture somewhat upright. Song a complex, sweet
warble learned by mimicry of other birds. Female sings during Breeds from Iberia and northwest Africa to western Siberia and
courtship. west China. Winters across sub-Saharan Africa.

FEEDING ECOLOGY AND DIET HABITAT

Gleans insects and spiders from low vegetation. Also takes Reedbeds and other emergent vegetation, sometimes in thickets
snails, and rarely berries. bordering water. Swamps, stream courses and savanna in winter.

REPRODUCTIVE BIOLOGY BEHAVIOR

Mostly monogamous, but serial monogamy and opportunistic Movements slow and clumsy. Song deep and slow, full of low
polygyny occur. Courtship may include aerial dance involving croaks and rattles. Male defends large territory.
both sexes. Nest is built by female; cylindrical cup of leaves
and plant material 3.3–6.6 ft (1–2 m) from ground in tall vege- FEEDING ECOLOGY AND DIET
tation, with rim woven around supporting vegetation. Three to
six eggs incubated and young, cared for by both parents, leave Insects, spiders, snails, small vertebrates (fish and frogs), some
nest after 11–12 days, remain dependent 15–19 days. fruits in non-breeding season. Forages in emergent vegetation,
trees and shrubs, and water surface.
CONSERVATION STATUS
REPRODUCTIVE BIOLOGY
Not threatened.
Many polygynous, but significant portion monogamous. Pair
SIGNIFICANCE TO HUMANS bond lasts only until nestling stage. Nest is built by female;
deep cylinder of reeds and leaves, suspended above water.
None known. ◆ Three to six eggs incubated 14 days by female; young fed by
both parents; fledglings dependent additional two weeks.
Great reed warbler
CONSERVATION STATUS
Acrocephalus arundinaceus
Not threatened.
SUBFAMILY
SIGNIFICANCE TO HUMANS
Sylviinae
None known. ◆
TAXONOMY
Icterine warbler
Acrocephalus arundinaceus Linnaeus, 1758. Subspecies griseldis
Hartlaub, 1891. Hippolais icterina

OTHER COMMON NAMES SUBFAMILY

French: Rousserolle turdoïde; German: Drosselrohrsänger; Sylviinae
Spanish: Carricero Tordal.
TAXONOMY
Acrocephalus arundinaceus
Hippolais icterina Viellot, 1817.
Breeding Nonbreeding
OTHER COMMON NAMES
Grzimek’s Animal Life Encyclopedia
French: Hypolaïs ictérine; German: Gelbspötter; Spanish:
Zarcero Icterino.

PHYSICAL CHARACTERISTICS

4.9–5.3 in (12.5–13.5 cm); 0.3–0.8 oz (8–23 g). Small warbler
with olive to gray-olive upperparts, yellowish underparts, pale
eye line, and long, thin bill.

DISTRIBUTION

France, Balkans, and Scandinavia east to southwest Siberia.
Winters in Southern Africa.

HABITAT

Open forests, orchards gardens and edges; often in lowlands
and river valleys.

BEHAVIOR

Active and vocal, but usually remains in foliage. Territorial on
breeding and wintering grounds. Song rapid, varied, including
musical phrases and mimicry.

17

Family: Old World warblers Vol. 11: Birds IV

Hippolais icterina

Breeding Nonbreeding

FEEDING ECOLOGY AND DIET Orthotomus sutorius

Insects, fruit in late summer. Forages in foliage, gleaning in- Resident
sects and sallying to catch aerial prey.
BEHAVIOR
REPRODUCTIVE BIOLOGY
Solitary or in pairs. Skulking but restless and active. Flicks tail
Monogamous. Nest is neat cup of plant material, decorated on side to side often. Song a loud, two-syllable phrase repeated in
outside with bark, paper, wool, etc., built by female in fork of series of three to four calls.
shrub or tree. Four to five eggs incubated by female for 13–15
days; young, cared for by both parents, leave nest after 12–16 FEEDING ECOLOGY AND DIET
days, independent after 1–2 weeks.
Insects and insect larvae, flower nectar. Forages on ground, in
CONSERVATION STATUS low vegetation and in trees.

Not threatened. REPRODUCTIVE BIOLOGY

SIGNIFICANCE TO HUMANS Nest is lined with spider silk, soft plant fibers, etc.; sewn by fe-
male from the leaves of a broad-leaved plant, usually well hid-
None known. ◆ den. Incubation of 3–5 eggs for 12 days and feeding nestlings
done by both parents.
Common tailorbird
CONSERVATION STATUS
Orthotomus sutorius
Not threatened.
SUBFAMILY
SIGNIFICANCE TO HUMANS
Sylviinae
None known. ◆
TAXONOMY
Yellow-bellied eremomela
Orthotomus sutoria Pennant, 1769.
Eremomela icteropygialis
OTHER COMMON NAMES
SUBFAMILY
English: Long-tailed tailorbird; French: Couterière à longe
queue; German: Rotstirn-Schneidervogel. Sylviinae

PHYSICAL CHARACTERISTICS TAXONOMY

5.1 in (13 cm); 0.2–0.4 oz (6–10 g). Small bird with long, thin, Eremomela icteropygialis Lafresnaye, 1822.
decurved bill. Back olive-green, underparts creamy, nape gray,
crown rufous, whitish eyeline. Short, spiky tail often held erect. OTHER COMMON NAMES

DISTRIBUTION English: Salvadori’s eremomela; French: Érémomèle à crou-
pion jaune; German: Gelbbauch-Eremomela; Spanish: Eremo-
Resident throughout India and Southeast Asia to Java and In- mela de Vientre Amarillo.
donesia.
PHYSICAL CHARACTERISTICS
HABITAT
3.9–4.3 in (10–11 cm); 0.26–0.33 oz (7.5–9.3 g). Small bird
Deciduous forest, scrubby clearings, gardens, scrubland, and with gray head, nape, back, and breast, yellow belly and under-
mangroves. tail. Dark eyeline surmounted by a lighter one; dark gray wing
and tail.
18
Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World warblers

Eremomela icteropygialis Sylvietta rufescens

Resident Resident

DISTRIBUTION OTHER COMMON NAMES

Widespread throughout non-forested sub-Saharan Africa. English: Cape crombec; French: Crombec à long bec; German:
Langschnabelsylvietta; Spanish: Sylvieta de Pico Largo.
HABITAT
PHYSICAL CHARACTERISTICS
Woodland, forest edge, scrub, gardens, and riparian areas.
3.9–4.7 in (10–12 cm); 0.3–0.8 oz (8–23 g). Small, nearly tail-
BEHAVIOR less bird with brownish gray upperparts, breast and flanks
buffy, throat and belly white. Black eyeline surmounted by a
Inconspicuous. Usually in pairs or groups of up to eight birds. light eyeline.
Territorial. Song is loud, twittery series of 7–8 notes. Mimicry
sometimes included. DISTRIBUTION

FEEDING ECOLOGY AND DIET Endemic resident in Southern Africa.

Foraging mostly in canopy and mid-level for small insects. HABITAT
Sometimes joins mixed-species foraging parties.
Thick brush, Acacia scrub, open and secondary woodland, often
REPRODUCTIVE BIOLOGY in drier areas.

Monogamous. Breeds primarily during monsoon season. Nest is BEHAVIOR
deep cup of plant down and spider silk, lined with grass, and sus-
pended from fork of a shrub or tree. One to three eggs are incu- Solitary or in pairs or family groups. Song a high, variable se-
bated 13–14 days by female; nestlings, fed by both parents, leave ries of trilled notes. Territorial. Flight bouncy.
after 15–16 days, fed by parents additional two or more weeks.
FEEDING ECOLOGY AND DIET
CONSERVATION STATUS
Insects, ticks and grass seeds. Forages methodically from bot-
Not threatened. tom to top of bushes and trees. Often joins mixed-species for-
aging parties.
SIGNIFICANCE TO HUMANS
REPRODUCTIVE BIOLOGY
None known. ◆
Monogamous. Nest is large, hanging bag of grasses, spider
Long-billed crombec webs, and plant fibers, close to ground and attached to the
lower limbs of a tree, often an acacia. One to three eggs incu-
Sylvietta rufescens bated for two weeks; nestlings fed by both parents for two
weeks before leaving.
SUBFAMILY
CONSERVATION STATUS
Sylviinae
Not threatened.
TAXONOMY
SIGNIFICANCE TO HUMANS
Sylvietta rufescens Vieillot, 1817.
None known. ◆
Grzimek’s Animal Life Encyclopedia
19

Family: Old World warblers Vol. 11: Birds IV

Chiffchaff nestlings cared by female; fledge after 14–16 days. Brood inde-
pendent after 10–19 days.
Phylloscopus collybita
CONSERVATION STATUS
SUBFAMILY
Not threatened.
Sylviinae
SIGNIFICANCE TO HUMANS
TAXONOMY
None known. ◆
Phylloscopus collybita Vieillot, 1817.
Arctic warbler
OTHER COMMON NAMES
Phylloscopus borealis
English: Common, Eurasian chiffchaff; French: Pouillot véloce;
German: Zilpzalp; Spanish: Mosquitero Común. SUBFAMILY

PHYSICAL CHARACTERISTICS Sylviinae

3.9–4.3 in (10–11 cm); 0.2–0.4 oz (6–12 g). Small warbler with TAXONOMY
short, fine bill, olive-green upperparts, white underparts,
brownish flanks, a dark line through the eye, surmounted by a Phyllopneuste borealis Blasius, 1858.
light line.
OTHER COMMON NAMES
DISTRIBUTION
French: Pouillot boréal; German: Wanderlaubsänger; Spanish:
Breeds from Iberia, British Isles and Scandinavia to Caspian Mosquitero Boreal.
Sea, northern Mongolia, eastern Siberia. Winters in Western
Europe, Mediterranean, northwest Africa, Middle East, India PHYSICAL CHARACTERISTICS
and sub-Saharan Africa.
4.1–5.1 in (10.5–13 cm); 0.3–0.5 oz (8–15 g). Medium-sized
HABITAT warbler, olive-green above, yellowish-white below, with a yel-
low wash in some plumages. Thin, clean whitish eye line; long
Open woodland with tall undergrowth, parks, and scrub, typi- wings with two whitish wing bars.
cally in lowlands.
DISTRIBUTION
BEHAVIOR
Breeds in boreal and subalpine zones from Scandinavia
Solitary or in pairs during breeding season. In small groups or throughout Asia to Japan and Western Alaska. Winters in
mixed flocks in migration and winter. Territorial. Song highly Southeast Asia, Wallacea.
variable; in nominate group it is series of two-note alternating
phrases. HABITAT

FEEDING ECOLOGY AND DIET Coniferous, deciduous, and mixed forest in taiga zone. Winters
in open woodlands, rainforest, forest edge, gardens, and man-
Forages from ground to canopy for insects, gleaning from fo- groves.
liage.
BEHAVIOR
REPRODUCTIVE BIOLOGY
Arboreal and active, with quick flight and habit of wing- and
Monogamous, sometimes facultatively polygynous. Nest is tail-flicking. Usually solitary or in pairs or small family groups.
built by the female; dome of dry gr–ss and other plant materi- Territorial; male often defends with song and wing-rattling
als with a side entrance, on or near the ground in thick vegeta- displays.
tion. Four to seven eggs, incubated 13–15 days by female;
FEEDING ECOLOGY AND DIET

Forages in foliage for insects and larvae, usually high, but occa-
sionally close to the ground.

Phylloscopus collybita Phylloscopus borealis

Resident Breeding Nonbreeding Breeding Nonbreeding

20 Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World warblers

REPRODUCTIVE BIOLOGY HABITAT

Monogamous, occasionally polygynous. Courtship involves Mid-level undergrowth of evergreen or mixed forest in high-
song, wing-rattling and wing-flapping displays. Nest is built by lands. Winters at lower elevations.
the female; dome of dry grass and other plant materials with a
side entrance on the ground in thick vegetation. Incubation of BEHAVIOR
5–7 eggs for 11–13 days by female; fledging takes 13–14 days,
young fed by both parents, brooded by female; young indepen- Usually solitary or in pairs. Rarely found in canopy. Song of
dent after two weeks. burkii is a loud, clear trill.

CONSERVATION STATUS FEEDING ECOLOGY AND DIET

Not threatened. Insects. Often joins mixed-species flocks in non-breeding season.

SIGNIFICANCE TO HUMANS REPRODUCTIVE BIOLOGY

None known. ◆ Little known. Nest is ball of mosses, grass, and other plant
fibers, lined with moss and lichen, and concealed along a bank
Golden-spectacled warbler or slope, often in tree roots. Four eggs are incubated by both
parents.
Seicercus burkii
CONSERVATION STATUS
SUBFAMILY
Not threatened.
Sylviinae
SIGNIFICANCE TO HUMANS
TAXONOMY
None known. ◆
Seicercus burkii Burton, 1836.
Little grassbird
OTHER COMMON NAMES
Megalurus gramineus
English: Yellow-eyed flycatcher warbler; French: Pouillot de
Burke; German: Goldbrillen-Laubsängerl Spanish: Curruca de SUBFAMILY
Burke.
Sylviinae
PHYSICAL CHARACTERISTICS
TAXONOMY
3.9–4.7 in (10–12 cm); 0.2–0.3 oz (6–9 g). Small, plump war-
bler, with bright green upperparts, bright yellow underparts, a Megalurus gramineus Gould, 1865.
short, broad tail with white undertail feathers. Crown streaked
black and gray, eyering yellow; yellow wingbar. OTHER COMMON NAMES

DISTRIBUTION English: Little marshbird; striated grassbird; little reedbird;
marsh warbler; French: Mégalure menue; German: Zwergschil-
Southern Asia, from India to south-central China to Thailand, fsänger.
Cambodia and Vietnam.
PHYSICAL CHARACTERISTICS

5.1–5.9 in (13–15 cm). Medium-sized warbler with brownish
upperparts, streaked dark; pale grayish underparts finely

Seicercus burkii Megalurus gramineus

Breeding Nonbreeding Resident

Grzimek’s Animal Life Encyclopedia 21

Family: Old World warblers Vol. 11: Birds IV

streaked with dark brown. Flight feathers graduated, edged PHYSICAL CHARACTERISTICS
with white.
5.5 in (14 cm); 0.5–0.7 oz (15–21 g). Medium-sized, with
DISTRIBUTION plumage ranging from slate gray in adult males to olive or
brown in females and juveniles. Crown is distinctive (black in
Southern and eastern Australia. adult males, rufous in females and juveniles). Wings are long
and pointed, with long primary projection. Bill is black, rela-
HABITAT tively long, and pointed. Legs long.

Dense vegetation of marshes, reedbeds, swamps, occasionally DISTRIBUTION
mangroves.
Breeds from British Isles and southern Scandinavia throughout
BEHAVIOR Western and Central Europe to coastal northwest Africa,
Mediterranean, Near East, and west to central Russia and
Solitary. Skulking. Song is three note, plaintive whistle. Flight northern Iran.
is weak. Some movement and nomadism occurs.
HABITAT
FEEDING ECOLOGY AND DIET
Forest with tall undergrowth, from riparian areas, parks and
Insects, insect larvae, spiders and other arthropods; also takes gardens to boreal forest and alpine forest to treeline.
aquatic mollusks.
BEHAVIOR
REPRODUCTIVE BIOLOGY
Arboreal and very active. Males territorial, defending with
Courtship involves chasing and wing-fluttering displays. Nest is song, displays, and agonistic behavior. Mimicry of other birds
suspended above water, a deep cup of grass and plant material, is occasionally incorporated into song. Mixed partial migrant:
lined with large feathers, and rimmed tops. Three to five eggs. individuals in northern range migrate south, while individuals
in southern range (the Mediterranean area) are residents or
CONSERVATION STATUS partial migrants.

Not threatened. FEEDING ECOLOGY AND DIET

SIGNIFICANCE TO HUMANS Feeds in trees and shrubs, gleaning insects and other arthro-
pods from leaves and branches. During migration and on win-
None known. ◆ tering grounds, fruits constitute a large part of the diet.

Blackcap REPRODUCTIVE BIOLOGY

Sylvia atricapilla Pairs solitary and territorial, generally monogamous. Site-fi-
delity is high in migratory populations. Courtship involves
SUBFAMILY construction, by the male, of several loose ‘cock nests’. The fe-
male completes the final nest, a fine cup typically located in
Sylviinae dense vegetation of a tree or shrub above ground. Both parents
incubate 2-6 eggs (typically 5) for 10–16 days. Feeding young
TAXONOMY in the nest (8–14 days) and after fledging (for about two weeks)
is also shared.
Sylvia atricapilla Linnaeus, 1758.
CONSERVATION STATUS
OTHER COMMON NAMES
Not threatened.
French: Fauvette à tête noire; German: Mönchsgrasmücke;
Spanish: Curraca Capirotada. SIGNIFICANCE TO HUMANS

A familiar songbird easily recognized by appearance and voice.
It is a model system for the study of the physiology and evolu-
tion of bird migration, and for the study of avian diet and en-
ergetics, especially as related to movement and seasonal food
availability. ◆

Sylvia atricapilla Whitethroat

Resident Breeding Nonbreeding Sylvia communis

22 SUBFAMILY

Sylviinae

TAXONOMY

Sylvia communis Latham, 1787.

OTHER COMMON NAMES

English: Common whitethroat, greater whitethroat; French:
Fauvette grisette; German: Dorngrasmücke; Spanish: Curruca
Zarcera.

Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World warblers

Sylvia communis DISTRIBUTION

Breeding Nonbreeding Breeds in Western and Central Europe, southern and coastal
Scandinavia, Turkey, North Africa, and Western and Central
PHYSICAL CHARACTERISTICS Siberia. Winters in sub-Saharan Africa.

5.5 in (14 cm); 0.4–0.9 oz (10–24 g). Medium-sized warbler HABITAT
with gray-brown upperparts, whitish underparts, buffy flanks,
gray (male) or brownish (female) cap and side of face, and a Open scrubland, farmland and forest edges, with mix of herba-
bright white throat. Folded wing shows significant rufous edg- ceous and low woody vegetation.
ing of coverts, secondaries and tertials. Long tail with white
outer tail feathers. BEHAVIOR

Typically solitary or in pairs. Skulking, but inquisitive. Male
song a short, scratchy warble. Territorial. Engages in song-
flight display, rising vertically from shrub, then swooping
back down.

FEEDING ECOLOGY AND DIET

Forages in low and mid-height vegetation, gleaning arthro-
pods.

REPRODUCTIVE BIOLOGY

Monogamous. Male builds ‘cock nests’. Nest a loose, deep cup
of grasses in dense, low, tangled vegetation. Three to six eggs
incubated 11–13 days, mostly by female; young brooded and
fed by both parents, leave nest after 10–12 days, remain with
parents 2–3 weeks.

CONSERVATION STATUS

Not threatened.

SIGNIFICANCE TO HUMANS

None known. ◆

Resources Periodicals
Alström, P. and U. Olsson. “The golden-spectacled warbler: a
Books
Ali, Salim, and S. Dillon Ripley. Handbook of the Birds of India complex of sibling species, including a previously
undescribed species.” Ibis 151 (1999): 545–568.
and Pakistan, 2nd ed. Vol. 8. Warblers to Redstarts. Delhi:
Oxford University Press, 1997. Barker, F. Keith, et al. “A phylogenetic hypothesis for
passerine birds: taxonomic and biogeographic implications
Baker, Kevin. Warblers of Europe, Asia and North Africa. of an analysis of nuclear DNA sequence data.” Proceedings of
Princeton: Princeton University Press, 1997. the Royal Society of London, Series B, 269 (2002): 295–308.

Cramp, Stanley, ed. The Birds of the Western Palearctic. Vol. 6. Catchpole, Clive K. “The evolution of mating systems in
Warblers. Oxford: Oxford University Press, 1992. Acrocephalus warblers.” Japanese Journal of Ornithology 44
(1995): 195–207.
Kaufman, Kenn. Lives of North American Birds. New York:
Houghton Mifflin, 1996. Helbig, Andreas J. and Ingrid Seibold. “Molecular phylogeny
of Palearctic-African Acrocephalus and Hippolais warblers
Parmenter, Tim and Clive Byers. A Guide to the Warblers of the (Aves: Sylviidae).” Molecular Phylogenetics and Evolution 11
Western Palearctic. Uxbridge, Middlesex: Bruce Coleman (1999): 246–260.
Books, 1991.
Irwin, Darren E., et al. “Speciation in a ring.” Nature 409
Shirihai, Hadoram, Gabriel Gargallo, and Andreas J. Helbig. (2001): 333–337.
Sylvia Warblers. Princeton: Princeton University Press,
2001. Leisler, B., et al. “Taxonomy and phylogeny of reed warblers
(genus Acrocephalus) based on mtDNA sequences and
Sibley, Charles G. and Jon E. Ahlquist. Phylogeny and morphology.” Journal fur Ornithologie 138 (1997): 469–496.
Classification of Birds. New Haven, Yale University Press,
1990. Matthew J. Sarver, BS

Urban, Emil K., Hilary C. Fry, and Stuart Keith. The Birds of
Africa. Vol. 5. San Diego: Academic Press, 1997.

Grzimek’s Animal Life Encyclopedia 23

▲

Old World flycatchers

(Muscicapidae)

Class Aves
Order Passeriformes
Suborder Passeri (Oscines)
Family Muscicapidae

Thumbnail description
A large, highly variable group consisting of about
135 species of songbirds. They are small-to
medium-sized perching birds, sometimes quite
colorful; most feed by making aerial sallies from
an exposed perch to catch their prey of flying
insects

Size
Body length 3–9 in (7.6–23 cm)

Number of genera, species
16 genera, 135 species, including 9 genera
(109 species) of Muscicapinae (typical Old
World flycatchers) and 7 genera (26 species) of
Platysteirinae (African flycatchers)

Habitat Distribution
Species occur in a wide range of habitats, Widespread in Eurasia and Africa with the greatest richness of species occurring in
including forest, woodland, savanna, grassland, tropical and subtropical Africa, India, and Southeast Asia
edges of waterbodies and wetlands, pasture
and other agricultural areas, and well-vegetated
gardens and residential areas

Conservation status
The World Conservation Union (IUCN) lists 18
species of the Muscicapidae as being at risk,
plus another 19 species as being Near
Threatened

Evolution and systematics Physical characteristics

As treated here, the Old World flycatchers (family Musc- The Old World flycatchers are a large and variable group
icapidae) include 16 genera and about 135 species, including consisting of about 135 species of songbirds. The subfamily
9 genera and 109 species of the subfamily Muscicapinae (the Muscicapinae includes small- to medium-sized birds, with a
typical Old World flycatchers) and 7 genera and 26 species body length generally ranging from 3–9 in (7.6–23 cm). The
of Platysteirinae (the African flycatchers). However, avian bill is relatively small, short, vertically (dorso-ventrally) flat-
systematists are actively studying the composition of this di- tened, and pointed. There are stout, well-developed bristles
verse group of birds and related ones. There is ongoing con- (known as rictal bristles) at the gape of the beak, which help
troversy about the relationship of the Muscicapidae to other increase the effectiveness at catching flying insects. The legs
designated families of passerine birds, and whether to com- are relatively short and the feet small but well suited to perch-
bine some of these with the Old World flycatchers. One ing. Coloration is highly variable among species, ranging from
classification joins the Muscicapidae with several related fam- dull gray or brown to bright blue or vermilion and sometimes
ilies, including the whistlers (Pachycephalidae), the fantails occurring in stark patterns. The sexes are colored alike in
(Rhipiduridae), and the monarchs (Monarchidae). That group many species, but are dimorphic in others. Young birds are
would include about 49 genera and 398 species. Some other usually relatively subdued in coloration and spotted on the
systematic treatments would include even more families than back and/or breast.
this, including the Old World warblers (Silviidae) and
thrushes (Turdidae). The tyrant flycatchers (Tyrannidae) are The subfamily Platysteirinae is also a varied group of
generally similar in appearance and behavior to the Musci- small- to medium-sized songbirds. The bill is small, short,
capidae, but this is because of convergent evolution as the vertically flattened, and slightly hooked at the tip. There are
families are not actually closely related. well-developed rictal bristles at the gape of the beak. The tail

Grzimek’s Animal Life Encyclopedia 25

Family: Old World flycatchers Vol. 11: Birds IV

subarctic regions and wintering in subtropical and tropical ar-
eas. During their migration they travel at night and feed dur-
ing the day. Species that breed in subtropical and tropical
regions are resident there, although some of them may un-
dertake seasonal movements between low- and high-altitude
habitats. Most species are non-gregarious, occurring only
singly, as pairs, or in small family groups consisting of par-
ents and their immature progeny. The Old World flycatch-
ers are strong and highly maneuverable fliers. Most species
feed by sitting on a conspicuous perch until a flying insect is
spotted, which is then pursued and caught in an aerial sally.
Some species also glean prey from foliage, bark surfaces, or
the ground. Some species have a well-developed song used to
proclaim and defend their breeding territory, but in others it
is relatively monotonous and weak. All species have calls used
to communicate with individuals of their species, or to alert
all nearby birds of the presence of a potential predator.

The colorful plumage of the blue-and-white flycatcher (Ficedula cyanome- Feeding ecology and diet
lana). (Photo by T. Shimba/VIREO. Reproduced by permission.)
Old World flycatchers use a variety of feeding tactics to
is short and the legs and feet are variable in length and stout- prey on insects and other arthropods. Most species use an ex-
ness. An area of bare skin around the eye is variable in color posed vantage point, such as a high perch in a tree, to spot
among species. The sexes are colored differently, with the flying insects, which are then caught in an aerial pursuit. Some
females being generally more subdued in pattern and hue. species also feed by gleaning prey from bark, branches, fo-
The plumage coloration of the male is mostly glossy black liage, or spider webs, or by swooping down to feed on arthro-
and white, while the female is usually more brown and some- pods spotted on the ground. Some species repeatedly return
times light rufous on the belly. In both the Muscicapinae and
Platysteirinae, relatively widespread species have been di-
vided into various gepgraphically distinct subspecies and
races that are distinct in elements of their size, plumage, be-
havior, and song.

Distribution

Species of typical Old World flycatchers are found widely
through Eurasia and Africa. However, the greatest richness
of species occurs in tropical Asia and Africa. Species of the
Platysteirinae occur only in Africa.

Habitat

Species of Old World flycatchers occur in a wide range of
habitats, including moist and dry forest, woodland, savanna,
grassland, edges of waterbodies and wetlands, pasture and
other agricultural areas, and well-vegetated gardens and res-
idential areas.

Behavior A cape puffback (Batis capensis) on its nest. (Photo by W. Tar-
boton/VIREO. Reproduced by permission.)
The northern species of Old World flycatchers are sea-
sonal migrants, breeding in higher-latitude temperate and Grzimek’s Animal Life Encyclopedia

26

Vol. 11: Birds IV Family: Old World flycatchers

to use a favorite perch for hunting, while others move about species as being Near Threatened. However, the conserva-
and frequently change their vantage point. tion status of many rare species in this group has not yet been
studied. Further research will undoubtedly add additional
Reproductive biology species to the listings. Examples of listed species at-risk in-
clude the Nimba flycatcher (Melaenornis annamarulae) of the
Old World flycatchers are highly territorial during their Ivory Coast and Guinea (Vulnerable), the streaky-breasted
breeding season, defending a nesting area from others of their jungle-flycatcher (Rhinomyias addita) of China (Near Threat-
species. They do this by proclaiming their territory by fre- ened), the white-browed jungle-flycatcher (Rhinomyias insig-
quent renditions of a song, and if this is not sufficient they nis) of the Philippines (Vulnerable), the Grand Comoro
will fight with intruders. They build a cup-shaped nest of flycatcher (Humblotia flavirostris) of the Comoro Islands (En-
grass, bark, and other plant fibers. The nest is generally placed dangered), the Lampobattang flycatcher (Ficedula bonthaina)
in the fork of a branch, on a ledge of a bank, or in a cavity in of Indonesia (Endangered), the red-tailed newtonia (Newto-
a tree, stump, or cliff. They lay two to seven spotted or mot- nia fanovanae) of Madagascar (Vulnerable), and the banded
tled eggs. In some species both parents participate in build- wattle-eye (Platysteira laticincta) of Cameroon (Endangered).
ing the nest and incubating the eggs, while in others only the All of these species are at-risk because of historical and on-
female does this. The incubation period ranges from about going habitat loss and fragmentation.
12–22 days. Both parents care for the nestlings and fledglings.
In some species, particularly of African flycatchers, immature Significance to humans
birds of previous nestings will help their parents raise a new
clutch of siblings. Old World flycatchers are not of direct importance to hu-
mans, other than the indirect economic benefits of ecotourism
Conservation status and bird-watching focused on seeing birds in natural habitats.
Moreover, it is crucial that research be undertaken to better
The World Conservation Union IUCN lists 18 species of understand the biology and habitat needs of the rare and en-
birds in the Muscicapidae as being at risk, plus another 19 dangered species of Old World flycatchers. In addition, their
critical habitats must be identified and rigorously protected
to prevent the extinction of these birds.

Grzimek’s Animal Life Encyclopedia 27

2
1

3

5
4

6

7

89

10
11

1. Silverbird (Empidornis semipartitus); 2. Black-and-white flycatcher (Bias musicus); 3. Abyssinian slaty flycatcher (Melaenornis chocolatinus); 4.
Orange-breasted blue flycatcher (Cyomis tickelliae); 5. Fraser’s forest-flycatcher (Fraseria ocreata); 6. Female shrike-flycatcher (Megabyas flam-
mulatus); 7. Black-throated wattle-eye (Platysteira peltata); 8. Cape batis (Batis capensis); 9. Female chestnut wattle-eye (Dyaphorophyia cas-
tanea) 10. Large-billed blue-flycatcher (Cyornis caerulatus); 11. Red-tailed newtonia (Newtonia fanovanae). (Illustration by Barbara Duperron)
28 Grzimek’s Animal Life Encyclopedia

2 3
1

5 7
4 6

9 10
8

1. Male collared flycatcher (Ficedula albicollis), spring plumage; 2. Dull-blue flycatcher (Eumyias sordida); 3. Male European pied flycatcher (Ficedula
hypoleuca), spring plumage; 4. Fulvous-chested jungle-flycatcher (Rhinomyias olivacea); 5. Female little slaty flycatcher (Ficedula basilanica) 6.
Brown-chested jungle-flycatcher (Rhinomyias brunneata); 7. Spotted flycatcher (Muscicapa striata); 8. Large niltava (Niltava grandis); 9. Ashy fly-
catcher (Muscicapa caerulescens); 10. Cassin’s flycatcher (Muscicapa cassini). (Illustration by Barbara Duperron)

Grzimek’s Animal Life Encyclopedia 29

Family: Old World flycatchers Vol. 11: Birds IV

Species accounts

Spotted flycatcher REPRODUCTIVE BIOLOGY

Muscicapa striata Builds a cup-shaped nest in a tree crotch, shallow tree-cavity,
or behind loose bark, and also uses nest-boxes. Lays four to six
SUBFAMILY greenish eggs with rust-colored spots.

Muscicapinae CONSERVATION STATUS

TAXONOMY Not threatened. A widespread and locally abundant species.

Muscicapa striata Pallas, 1764. SIGNIFICANCE TO HUMANS

OTHER COMMON NAMES None known, except for the economic benefits of bird-watch-
ing. ◆
French: Gobemouche gris; German: Grauschnäpper; Spanish:
Papamoscas Gris. Ashy flycatcher

PHYSICAL CHARACTERISTICS Muscicapa caerulescens

The body length is about 5 in (13–14 cm). Both sexes are col- SUBFAMILY
ored alike, having a brownish gray back, head, and tail and a
white belly and throat streaked with gray. The juvenile is more Muscicapinae
spotted. Various subspecies have been described based on
plumage and song characters. TAXONOMY

DISTRIBUTION Muscicapa caerulescens Hartlaub, 1865.

Breeds widely in northern and central Europe and European OTHER COMMON NAMES
and central Russia, and winters in southern Africa.
English: Ashy alseonax, blue-gray flycatcher, cinereus fly-
HABITAT catcher; French: Gobemouche à lunettes; German: Schiefer-
schnäpper; Spanish: Papamoscas Ahumado.
Breeds in temperate forest edges, woods, parks, orchards, and
gardens. PHYSICAL CHARACTERISTICS

BEHAVIOR The body length is 5.5–6 in (14–16 cm). The sexes are colored
similarly, with a light blue-gray back, gray-white underparts, a
A migratory species. Pairs of breeding birds defend a territory.
Winters as single birds. Often flicks its wings and tail when
perched. The song is delivered from a prominent perch, and is
a series of about six squeaky notes.

FEEDING ECOLOGY AND DIET

Spots flying insects from a prominent perch and then pursues
the prey by an aerial sally. Usually returns persistently to the
same perch.

Muscicapa striata Muscicapa caerulescens

Breeding Nonbreeding Resident

30 Grzimek’s Animal Life Encyclopedia

Vol. 11: Birds IV Family: Old World flycatchers

white eye-ring, and a black line bordered by a white one ex- Muscicapa cassini
tending through the eye. Several subspecies have been identi-
fied, which differ somewhat in coloration. Resident

DISTRIBUTION FEEDING ECOLOGY AND DIET

A resident species of much of southern Africa. An active hunter that searches for flying insects from an ex-
posed perch, such as a stump or dead tree in the water or from
HABITAT an overhanging branch. Usually returns to its original perch af-
ter each sally. Sometimes swoops to take prey from a spider
Occurs in a wide range of tropical forest-edges and moist open web or the ground.
woodlands and savannas, and also cleared and agricultural ar-
eas. Does not occur in the interior of closed forest. Occurs as REPRODUCTIVE BIOLOGY
high as 5,900 ft (1,800 m).
Builds a cup-shaped nest of grass and other fibers. The nest is
BEHAVIOR placed close to the ground in a shallow cavity in a stump, in
other kinds of tree-crevices, or at a narrow branch-fork. Lays
A nonmigratory species. Pairs of breeding birds defend a terri- two light-green, finely speckled eggs.
tory. The song consists of three to five high-pitched notes.
CONSERVATION STATUS
FEEDING ECOLOGY AND DIET
Not threatened. A widespread and locally abundant species.
Hunts from a perch for flying insects in the upper canopy.
Usually returns to its original perch after an aerial sally. SIGNIFICANCE TO HUMANS

REPRODUCTIVE BIOLOGY None known, except for the economic benefits of bird-
watching. ◆
Builds a bulky cup-shaped nest of moss, grass, and rootlets
lined with finer material. Nest is placed in a shallow tree- Pied flycatcher
cavity, a bark crevice, or at a narrow branch-fork. Lays two or
three creamy colored, finely speckled eggs. Ficedula hypoleuca

CONSERVATION STATUS SUBFAMILY

Not threatened. A widespread and locally abundant species. Muscicapinae

SIGNIFICANCE TO HUMANS TAXONOMY

None known, except for the economic benefits of bird-watch- Ficedula hypoleuca Pallas, 1764.
ing. ◆
OTHER COMMON NAMES
Cassin’s flycatcher
English: European pied flycatcher; French: Gobemouche noir;
Muscicapa cassini German: Trauerschnäpper; Spanish: Papamoscas Cerrojillo.

SUBFAMILY 31

Muscicapinae

TAXONOMY

Muscicapa cassini Heine, 1859.

OTHER COMMON NAMES

English: Cassin’s alseonax, Cassin’s gray flycatcher; French:
Gobemouche de Cassin; German: Cassinschnäpper; Spanish:
Papamoscas de Cassin.

PHYSICAL CHARACTERISTICS

The body length is about 5.5 in (14 cm). The sexes are colored
similarly, with a bluish gray back, black wings and tail, and
white underparts with gray flanks and chest.

DISTRIBUTION

A resident species of much of western tropical Africa.

HABITAT

Occurs in the vicinity of rivers, streams, and other surface wa-
ters within humid, lowland, tropical forest. Occurs as high as
about 5,900 ft (1,800 m).

BEHAVIOR

A nonmigratory species. Pairs of breeding birds defend a linear
territory along a watercourse, or a wider one in flooded forest.
The song consists of a medley of whistles, buzzes, and chirps
sung in bouts of several minutes.

Grzimek’s Animal Life Encyclopedia

Family: Old World flycatchers Vol. 11: Birds IV

Ficedula hypoleuca

Breeding Nonbreeding

PHYSICAL CHARACTERISTICS Ficedula albicollis

The body length is 5 in (13–14 cm). The male is colored strik- Breeding Nonbreeding
ingly black-and-white, with a black back and head, a white
belly and throat, and white wing-flashes. The female and juve- OTHER COMMON NAMES
nile are gray above, with a white belly and throat, and white
wing-flashes. The winter male is colored more grayish. English: Half-collared flycatcher; semicollared flycatcher;
French: Gobemouche à collier; German: Halsbandschnäpper;
DISTRIBUTION Spanish: Papamoscas Collarino.

Breeds widely in northern and central Europe and European PHYSICAL CHARACTERISTICS
Russia, and winters in western equatorial Africa.
The body length is about 4.5 in (11.5 cm), with the male col-
HABITAT ored black-and-white, with a black back and head, an interven-
ing white collar at the nape, a white belly and throat, and
Breeds in temperate forest, woods, parks, orchards, and gardens. white wing-flashes. The female and juvenile are gray with a
white belly and throat and white wing-flashes. The winter male
BEHAVIOR is colored more grayish. Various subspecies have been de-
scribed based on plumage and song characters.
A migratory species. Pairs of breeding birds defend a territory.
Winters as single birds. Often raises its fanned tail while DISTRIBUTION
perched. The song is delivered from a prominent perch, and is
a varied series of loud trills. Breeds widely in Europe, and winters in eastern equatorial
Africa.
FEEDING ECOLOGY AND DIET
HABITAT
Catches flying insects by an aerial sally from a prominent
perch. Usually returns persistently to the same perch. Breeds in temperate forest, woods, parks, orchards, and gardens.

REPRODUCTIVE BIOLOGY BEHAVIOR

Nests in a tree-cavity, and also uses nest-boxes. Lays five to A migratory species. Pairs of breeding birds defend a territory.
seven blue eggs. Winters as single birds. Often raises its fanned tail while
perched. The song is delivered from a prominent perch, and is
CONSERVATION STATUS a varied series of soft trills.

Not threatened. A widespread and locally abundant species. FEEDING ECOLOGY AND DIET

SIGNIFICANCE TO HUMANS Catches flying insects by an aerial sally from a prominent
perch. Usually returns persistently to the same perch.
None known, except for the economic benefits of bird-
watching. ◆ REPRODUCTIVE BIOLOGY

Collared flycatcher Nests in a tree-cavity or holes in rock walls, and also uses nest-
boxes. Lays five to seven blue eggs.
Ficedula albicollis
CONSERVATION STATUS
SUBFAMILY
Not threatened. A widespread and locally abundant species.
Muscicapinae
Grzimek’s Animal Life Encyclopedia
TAXONOMY

Ficedula albicollis Temminck, 1795.

32

Vol. 11: Birds IV Family: Old World flycatchers

SIGNIFICANCE TO HUMANS BEHAVIOR

None known, except for the economic benefits of bird- A nonmigratory species. Pairs of breeding birds defend a terri-
watching. ◆ tory. A quiet, skulking species of the forest understory. The
song is a series of warbling phrases.
Little slaty flycatcher
FEEDING ECOLOGY AND DIET
Ficedula basilanica
Searches from a low perch for flying insects.
SUBFAMILY
REPRODUCTIVE BIOLOGY
Muscicapinae
Builds a cup-shaped nest.
TAXONOMY
CONSERVATION STATUS
Ficedula basilanica Sharpe, 1877.
Vulnerable. A rare and declining species. Much of its original
OTHER COMMON NAMES natural habitat has been lost to the development of subsistence
agriculture, commercial plantations, logging, mining, and in
French: Gobemouche de Basilan; German: Schiefergrund- one place, a golf course. Some of its breeding habitat occurs in
schnäpper; Spanish: Papamoscas Pizarro Chico. protected areas, but is not necessarily safe from disturbance
there. Areas of its critical breeding habitat must be protected.
PHYSICAL CHARACTERISTICS
SIGNIFICANCE TO HUMANS
The body length is about 5 in (12 cm). The male is colored
slate-gray on the back, head, and tail and has a white belly. None known, except for the economic benefits of bird-
The female is more brownish. watching. ◆

DISTRIBUTION Brown-chested jungle-flycatcher

Vulnerable. A rare, endemic, nonmigratory species that only Rhinomyias brunneata
occurs on the islands of Samar, Leyte, Dinagat, Mindanao, and
Basilan in the Philippine archipelago. SUBFAMILY

HABITAT Muscicapinae

Occurs in lowland, humid, evergreen, primary and mature sec- TAXONOMY
ondary tropical forest, including selectively logged areas. It se-
lects microhabitat within the dense forest understorey. It Rhinomyias brunneata Slater, 1897.
occurs over the altitudinal range of sea level to 3,900 ft
(0–1,200 m). OTHER COMMON NAMES

English: Brown-chested flycatcher, Chinese olive flycatcher;
French: Gobemouche à poitrine brune; German: Weisskehl-
Dschungelschnäpper; Spanish: Papamoscas Selvático de Gar-
gantilla Blanca.

Ficedula basilanica Rhinomyias brunneata

Resident Breeding Nonbreeding

Grzimek’s Animal Life Encyclopedia 33