Comparative Anatomy of the External and Middle Ear of Palaeognathous Birds

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Editors F. Beck, Melbourne W. Hild, Galveston W. Kriz, Heidelberg J.E. Pauly, Little Rock Y. Sano, Kyoto T.H. Schiebler, Wiirzburg

Springer Berlin Heidelberg New York Barcelona Budapest Hong Kong London Milan Paris Tokyo

J. Matthias Starck

Comparative Anatomy of the External and Middle Ear of Palaeognatholls Birds

With 57 Figures and 4 Tables

Springer

Dr. J. Matthias Starck Eberhard-Karls-U niversitat Tiibingen Zoologisches Institut Auf der Morgenstelle 28 72076 Tiibingen, Germany

ISBN-13: 978-3-540-58991-4 e-ISBN-13: 978-3-642-79592-3 DOl: 10.1007/ 978-3-642-79592-3 Library of Congress Cataloging·in-Publication Data. Starck, J. Matthias, 1958Comparative anatomy of the external and middle ear of palaeognathous birdslJ. Matthias Starck. p. cm.-(Advances in anatomy, embryology, and cell biology; v. 131) Includes bibliographical references (p. ) and index. ISBN-13: 978-3-540-58991-4. Ratites-Anatomy. 2. Ear-Anatomy. I. Title. II. Series. QL801.E67 v. 131 [QL697] 574.4 s-dc20 [598.5'0448] 95-8260

© Springer-Verlag Berlin Heidelberg 1995 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Product liability: The publishers cannot guarantee the accuracy of any information about dosage and application contained in this book. In every individual case the user must check such information by consulting the relevant literature. Typesetting: Best-set Typesetter Ltd., Hong Kong SPIN: 10474714

27/3136/SPS - 5 4 3 2 1 0 - Printed on acid-free paper

Contents

1

Introduction, ............................ .

1

1.1 1.2

Objectives of this Study ................... . Literature Review ........................ .

1 2

1.3

Theoretical Concepts: Questions and "Answers" .......................... . Description and Representation ............ . Comparative Approach, Homology, Systematics, and Phylogeny ............... . Functional Approach ..................... . Ecomorphology .......................... .

1.3.1 1.3.2 1.3.3 1.3.4 1.4 1.4.1 1.4.2 1.4.3 1.4.4 1.5 1.5.1 1.5.1.1 1.5.1.2 1.5.1.3 1.5.1.4 1.5.2

Basic Functional Concepts of the Avian Middle Ear: The Transfer Function .................... . Area Ratio .............................. . Lever Systems ........................... . Curved Membrane and Resonance Effects ... . Impedance of the Middle Ear .............. . Evolutionary History of Middle Ear Structures .................. . Homologies of the Avian Middle Ear and the Hyoid Arch ...................... . Stapes .................................. . Tympanic Process of the Extracolumella .... . Processus supracolumellaris ............... . Processus infracolumellaris ................ . Phylogenetic Relationship of Palaeognathous and Basic Neognathous Birds .............. .

4

5 5 6 7

7

8 8 9 9 10 10 10 11 11 12

13

2

Materials and Methods . ................... .

17

2.1 2.2 2.3 2.4

Species List .