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The Inner Bird kaiser1.P65 1 1/29/2007, 9:18 PM kaiser1.P65 2 1/29/2007, 9:18 PM GARY W. KAISER The Inner Bird Anatomy and Evolution kaiser1.P65 3 1/29/2007, 9:18 PM © UBC Press 2007 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without prior written permission of the publisher, or, in Canada, in the case of photocopying or other reprographic copying, a licence from Access Copyright (Canadian Copyright Licensing Agency), www.accesscopyright.ca. 15 14 13 12 11 10 09 08 07 5 4 3 2 1 Printed in Canada on ancient-forest free paper (100% post consumer recycled) that is processed chlorine- and acid-free, with vegetable-based inks. Library and Archives Canada Cataloguing in Publication Kaiser, Gary W. The inner bird: anatomy and evolution / Gary W. Kaiser. Includes bibliographical references and index. ISBN 978-0-7748-1343-3 1. Birds – Anatomy. 2. Birds – Evolution. I. Title. QL697.K24 2007 571.3’18 C2007-900240-4 UBC Press gratefully acknowledges the financial support for our publishing program of the Government of Canada through the Book Publishing Industry Development Program (BPIDP), and of the Canada Council for the Arts, and the British Columbia Arts Council. Printed and bound in Canada by Friesens Set in Arrus and Aurora by Artegraphica Design Co. Ltd. Text designer: Irma Rodriguez Project editor: Francis Chow UBC Press The University of British Columbia 2029 West Mall Vancouver, BC V6T 1Z2 604-822-5959 / Fax: 604-822-6083 www.ubcpress.ca kaiser1.P65 4 1/29/2007, 9:18 PM For Dr. Alice Wilson, Dr. Donald C. Maddox, Mr. Herb Groh, and all the other adult mentors of the Macoun Field Club, the Ottawa Field-Naturalists’ program for young people kaiser1.P65 5 1/29/2007, 9:18 PM kaiser1.P65 6 1/29/2007, 9:18 PM Contents Figures and Tables / ix Acknowledgments / xiii Introduction / 1 Part 1: What Is a Bird? 1 The Bird beneath the Feathers / 15 How adaptations for flight allow a bird to earn a living on the ground 2 A Bird Is an Animal with Hollow Bones / 40 Special features of avian bones and their assembly as a skeleton 3 A Bird Is Like a Dinosaur / 86 Features shared by birds and their reptilian ancestors 4 A Bird Is Not So Like a Dinosaur / 108 Features that distinguish birds from their reptilian ancestors Part 2: What Kind of Bird Is It? 5 The Kinds of Birds / 132 Classification and taxonomy 6 That Bird Is Different from the Other One / 168 Evolutionary relationships among families of birds kaiser1.P65 7 1/29/2007, 9:18 PM Part 3: How Does a Bird Fly? 7 Feathers and Feathered Dinosaurs / 205 Flight in early birds and birdlike animals 8 Birds with a Modern Shape / 235 Birds cease to look like feathered dinosaurs and become skilled fliers 9 Birds on Land / 275 A robust design helps birds compete with mammals 10 Birds at Sea / 305 Sophisticated aerodynamics take birds onto oceans Conclusion / 344 Appendices 1 Birds in Relation to Other Vertebrate Animals / 346 2 Geological Time Scale / 347 Glossary of Ornithological Terms / 349 Literature Cited / 360 Index / 372 kaiser1.P65 8 1/29/2007, 9:18 PM Figures and Tables Figures 1.1 Major elements in the avian skeleton / 18 1.2 Dorsal view of the skull of a Pacific Loon / 19 1.3 The flattened disc of the avian eye in cross-section / 22 1.4 The specialized eyes of an owl / 23 1.5 The hyoid apparatus of the Chattering Lory, a parrot / 26 1.6 Relationship of the brain and eye in the skull of a Yellow-billed Hornbill / 28 1.7 Caudal or posterior views of the skulls in four birds, showing the positions of the foramen magnum and occipital condyle / 29 1.8 Two extremes in the shapes of the tarsometatarsus / 32 1.9 The basic arrangements of toes in birds / 34 1.10 Variations in the proportions of the major skeletal elements in the wings of three aerial specialists among small forest birds / 36 2.1 Sagittal section through the upper humerus of a Common Murre / 49 2.2 Sagittal section through the skull of a Helmeted Hornbill / 49 2.3 Movement of the upper jaw in parrots / 58 2.4 Neck vertebrae supported by a sling of neck ligaments in a chicken / 61 2.5 Two different methods of lowering the head in extremely long-necked birds / 62 2.6 The first two cervical vertebrae in a Brandt’s Cormorant / 63 2.7 Thoracic vertebra from a Common Loon / 68 2.8 Skeleton of the thoracic and sacral regions in the Hoatzin and Common Murre / 70 2.9 Lateral and ventral views of vertebrae in the Cassin’s Auklet near the meeting of the thoracic and sacral regions / 72 2.10 The lumbar interface in a Double-crested Cormorant / 73 2.11 The lumbar interface in a Turkey Vulture and a Great Horned Owl / 74 2.12 Cross-section through the hipbones of a Bald Eagle / 75 2.13 The hypotarsus in a Common Raven / 84 kaiser1.P65 9 1/29/2007, 9:18 PM x Figures and Tables 3.1 Squamosal and parasagittal stances in the hind limbs of primitive vertebrates / 87 3.2 The sagittal plane and directional orientation in a bird / 88 3.2 Nineteenth-century netsuke capturing Richard Owen’s reconstruction of an iguanodon / 105 4.1 Expansion of the hipbones in a Belted Kingfisher to create an “abdominal vault” over the intestines and reproductive system / 115 4.2 Relative size of the hipbones and an egg in the Western Grebe and the Rhinoceros Auklet / 117 4.3 The relationship between the distribution of the heavy digestive organs and the angle of the pubic bone in four members of the archosaur lineage / 118 5.1 Pierre Belon’s classic comparison of human and bird skeletons, published in 1555, shortly after the first publication of Aristotle in Latin / 136 6.1 A family tree for the major groups of birds based on a 1981 cladistic analysis by Joel Cracraft / 170 6.2 The family tree of living birds derived by Sibley and Ahlquist from the results of biomolecular analyses / 181 6.3 The phylogeny of birds based on variation in the b-fibrinogen gene from nuclear DNA / 197 7.1 The distribution of protofeathers and modern-looking feathers in a genealogical relationship for advanced theropod dinosaurs / 220 7.2 The forelimb skeleton superimposed on a recent reconstruction of Archaeopteryx / 229 7.3 Silhouettes of a Great Hornbill in flight / 232 7.4 Calculating the aspect ratio of a bird’s wings / 232 8.1 Three typical fossils of Confuciusornis sanctus, showing the large hands and feet and the massive humerus with the hole near the shoulder / 238 8.2 Fossil B 065 from the Wenya Museum, Jinzhou City, China / 239 8.3 A reconstruction of Confuciusornis in flight / 240 8.4 The pectoral girdle of Confuciusornis sanctus / 243 8.5 A comparison of the pelvic girdles in Confuciusornis sanctus and the Turkey Vulture / 244 8.6 Liaoxiornis delicatus, a modern type of bird from the Cretaceous, with a well-developed furcula / 250 8.7 The family tree for birds developed by Julia Clarke after her analysis of the Ichthyornis fossils / 256 8.8 An abbreviated version of L.M. Chiappe’s phylogeny for early birds, showing the groups included in the Pygostylia / 257 8.9 Reconstruction of the sternum and furcula from a member of the Enantiornithes (ball-shouldered birds) (Cathayornis) and the same bones from a late embryo of the Hoatzin / 263 kaiser1.P65 10 1/29/2007, 9:18 PM Figures and Tables xi 8.10 Diagrammatic cross-sections showing the types of construction used in the coracoid bone / 264 8.11 The furcula in four fossil birds / 266 8.12 The furcula in two modern birds / 267 9.1 The global distribution of weights among species in three of the most numerous groups of forest birds / 283 9.2 Brachial indices for various groups of living birds / 289 9.3 A chickadee rolls its wings into a tube at the beginning of the recovery stoke / 291 9.4 The division of flying birds into three groups according their wing shape and weight / 296 9.5 The range of variation in weight and aspect ratio for the Charadriiformes and the Procellariiformes / 297 9.6 Increases in average weight among clades of flying birds in the sequence presented in Sibley and Ahlquist’s biomolecular phylogeny / 299 9.7 Increases in average wing loading among clades of flying birds in the sequence presented in Sibley and Ahlquist’s biomolecular phylogeny / 299 9.8 Increases in average aspect ratio among clades of flying birds in the sequence presented in Sibley and Ahlquist’s biomolecular phylogeny / 300 9.9 A dendrogram derived from a phylogeny of birds posted on a “Vertebrate Notes” website that has since been closed / 301 10.1 Comparison of wing skeletons in five wing-propelled diving birds and a non-diving gull / 315 10.2 Outline of an auk wing, partially folded for underwater flight / 317 10.3 Thoracic skeleton of a frigatebird, showing the very large furcula and coracoids fused to each other and to the scapulas / 319 10.4 A mathematical expression of the power required for aerial flight, developed by J.M.V. Rayner / 323 10.5 The patagial ossicle that appears in the wings of albatrosses and some of their relatives / 329 10.6 Relative egg size compared with wing loading in two important groups of seabirds / 332 10.7 The weight of food loads delivered to nestlings compared with wing loading for two important groups of seabirds / 333 Tables 2.1 Hans Gadow’s allocation of vertebrae to zones in the avian spinal column / 66 5.1 Hans Gadow’s 1893 classification of birds / 147 6.1 Proposed names of clades to help resolve confusion around the use of the term “Aves” as it applies to both fossil animals and living birds / 174 6.2 The clades of living birds and their immediate relatives that survived the end of the Cretaceous / 175
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