Vertebrate Flight: Mechanics, Physiology, Morphology, Ecology and Evolution

303 Pages·1990·8.604 MB·English

by  Dr. Ulla M. Norberg (auth.)

Zoophysiology Volume 27
Editors
S.D. Bradshaw W Burggren
H.C. Heller S. Ishii H. Langer
G. Neuweiler D. J. Randall
Zoophysiology
Volumes already published in the series:
Volume 1: P.I. Bentley Volume 15: T. Mann
Spermatophores (1984)
Endocrines and Osmoregulation
(1971) Volume 16: P. Bouverot
Volume 2: L. Irving Adaption to Altitude-Hypoxia
Arctic Life of Birds and Mammals in Vertebrates (1985)
Including Man (1972)
Volume 17: R.J. R Smith
Volume 3: A. E. Needham The Control of Fish Migration (1985)
The Significance of Zoochromes (1974)
Volume 18: E. Gwinner
Volume 4/5: A. C. Neville Circannual Rhythms (1986)
Biology of the Arthropod Cuticle
(1975) Volume 19: I. C. Ruegg
Calcium in Muscle Activation (1986)
Volume 6: K. Schmidt-Koenig
Migration and Homing in Animals Volume 20: l.-R. Truchot
(1975) Comparative Aspects of Extracellular
Acid-Base Balance (1987)
Volume 7: E. Curio
The Ethology of-Predation (1976) Volume 21: A. Epple and I. E. Brinn
The Comparative Physiology of the
Volume 8: W. Leuthold
Pancreatic Islets (1987)
African Ungulates (1977)
Volume 9: E. B. Edney Volume 22: W.H. Dantzler
Water Balance in Land Arthropods Comparative Physiology of the
(1977) Vertebrate Kidney (1988)
Volume 10: H.-U. Thiele Volume 23: G. L. Kooyman
Carabid Beetles in Their Environments Diverse Divers (1989)
(1977)
Volume 24: S. S. Guraya
Volume 11: M.H.A. Keenleyside Ovarian Follicles in Reptiles and
Diversity and Adaptation in Birds (1989)
Fish Behaviour (1979)
Volume 25: G.D. Pollak and
Volume 12: E. Skadhauge I. H. Casseday
Osmoregulation in Birds (1981) The Neural Basis of Echolocation in
Bats (1989)
Volume 13: S. Nilsson
Autonomic Nerve Function in the Volume 26: G.A. Manley
Vertebrates (1983) Peripheral Hearing Mechanisms in
Reptiles and Birds (1989)
Volume 14:A.D. Hasler
Olfactory Imprinting and Homing Volume 27: U. M. Norberg
in Salmon (1983) Vertebrate Flight (1989)
V.M. Norberg
Vertebrate Flight
Mechanics, Physiology, Morphology,
Ecology and Evolution
With 103 Figures
Springer-Verlag Berlin Heidelberg New York
London Paris Tokyo Hong Kong
Dr. ULLA M. NORBERG
Department of Zoology
Division of Zoomorphology
University of Goteborg
Box 25059
S-400 31 Goteborg
ISBN-13:978-3-642-83850-7 e-ISBN-13:978-3-642-83848-4
DOl: 10.1007/978-3-642-83848-4
This work is subject to copyright. All rights are reserved, whether the whole or part of
the material is concerned, specifically the rights of translation, reprinting, re-use of
illustrations, recitation, broadcasting, reproduction on microfilms orin other ways, and
storage in data banks. Duplication of this publication or parts thereof is only permitted
under the provisions of the German Copyright Law of September 9, 1965, in its version
of June 24, 1985, and a copyright fee must always be paid. Violations fall under the
prosecution act of the German Copyright Law.
© Springer-Verlag Berlin Heidelberg 1990
Softcover reprint of the hardcover 1st edition 1990
The use of 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.
Typesetting: International Typesetters Inc., Makati, Philippines
213113145-543210 - Printed on acid-free paper
Dedicated to Ake, Peter, and Bjorn
Preface
It has been great fun to write this book, even though it has taken longer than
planned, and occasionally been exasperating. The most difficult problem was
deciding what to exclude among so many interesting things, because the
available material usually exceeded the space. Because a book like this covers
so many aspects, each component must be limited.
This book is intended for graduate and undergraduate students as well as
professional scientists who want to work with animal flight or to gain some
insight into flight mechanics, aerodynamics, energetics, physiology, morpho
logy, ecology and evolution. My aim has not been to give the whole mathe
matical explanation of flight, but to provide an outline and summary of the
main theories for the understanding of how aerofoils respond to an airflow. I
also hope to give the reader some insight into how flight morphology and the
various wing shapes have evolved and are adapted to different ecological
niches and habitats.
I would never have written or completed this book without the encourage
ment and help of my family and my friends. lowe tremendous thanks to my
husband Ake Norberg for endless discussions on various problems within
flight mechanics, morphology, ecology and evolution, and for reading and
commenting on the manuscript, and to my sons Peter and Bjorn (born 1974 and
1977) for patience and encouragement rather than complaints when my time
with them was limited. I also thank Peter for checking the references. I am
most grateful to M. B. Fenton, in particular, for reading and commenting on
the whole manuscript, and to C.J. Pennycuick, J.M.V. Rayner, H.D.J.N.
Aldridge, P. J. Butler, G. Goldspink, and F. R. Hainsworth for reviewing
various parts of it. Special thanks go to G. R. Spedding for allowing me to use
his exellent stereo-photographs ofthe vortex wake of a pigeon. I am indebted
to the series editor, G. Neuweiler, for asking me to write the book and for his
patience, and to various colleagues who have encouraged the writing by asking
questions like "When will the book be ready?". I thank Academic Press,
Harvard University Press, The Company of Biologists Limited, The Royal
Society, The University of Chicago Press, The Zoological Society of London
and Springer-Verlag for permission to reproduce figures from their journals.
The Swedish Natural Science Research Council has generously supported my
research, much of which has been included in various sections ofthe book.
It is impossible to write a book without any errors. Although the manu
script has been checked by specialists, the book may still contain several
mistakes which are my responsibility.
Askim, Summer 1989 ULLA M. NORBERG
VII
Contents
Chapter 1. Introduction 1
1.1 Gliders 2
1.2 Active Flyers . 4
1.3 Outline ofthe Book 6
Chapter 2. Basic Aerodynamics 8
2.1 Introduction . . . . . . 8
2.2 The Flow Around an Aerofoil . 13
2.2.1 Bernoulli's Equation 13
2.2.2 Reynolds Number . . . . . . . 16
2.2.3 Boundary Layer . . . . . . . . 17
2.3 Blade-Element and Momentum Jet Theories 19
2.3.1 Lift and Drag . . . . . 20
2.3.2 Power Required to Fly ...... . 24
2.4 Vortex Theory of Flight . . . . . . . 30
2.4.1 Bound, Trailing and Starting Vortices 30
2.4.2 Steady Motion . . . . . . . 32
2.4.3 Lifting-Line Theories ... 33
2.4.4 Quasi-Steady Assumption . 36
2.4.5 Unsteady Effects ..... 37
Chapter 3. Physiology of Flight 40
3.1 Introduction ..... . 40
3.2 Energy and Mechanical Efficiency 40
3.3 Metabolic Rates ... . 42
3.4 Oxygen Uptake ..... . 44
3.4.1 Respiratory Mechanics .. 44
3.4.2 Respiratory Gas Exchange 45
3.4.3 Circulation ....... . 48
3.4.4 Oxygen Consumption Versus Body Mass 50
3.4.5 Mass Loss . . . . . . . . 51
3.4.6 Heat Loss and Exchange ....... . 52
IX
3.5 Altitudinal Changes . . . . . . . . . . . . . . . . . . . 54
3.6 Estimates for Cost of Flight . . . . . . . . . . . . . . . 55
3.6.1 Direct Measurements of O Uptake and CO Production 55
2 2
3.6.2 Mass Loss ........... . 57
3.6.3 Doubly Labelled Water Method . . . . . . . . . . . 57
3.6.4 Radio Telemetry. . . . . . . . . . . . . . . . . . . 58
3.7 Flight Duration, Flight Range and Cost of Transport 59
Chapter 4. Morphological Flight Parameters 61
4.1 Introduction...... 61
4.2 Lenghts, Areas, Masses 61
4.3 Wing Shape . . . . . . 63
4.4 Weis-Fogh's and Ellington's Shape Parameters 63
Chapter 5. Gliding Flight 65
5.1 Introduction... 65
5.2 Gliding Performance 65
5.3 Effects of Change in Wingspan on Gliding Performance 70
5.4 Lifting Line Theory . . . . . . . . . 70
5.5 Flap-Gliding ............ . 71
5.6 Stability and Control of Movements. 73
5.6.1 Pitch. 73
5.6.2 Roll 74
5.6.3 Yaw . 75
Chapter 6. Soaring 76
6.1 Introduction 76
6.2 Soaring Methods 76
6.2.1 Slope Soaring .. 76
6.2.2 Thermal Soaring . 77
6.2.3 Gust, Frontal and Wave Soaring 78
6.2.4 Dynamic Soaring ........ 78
6.3 Soaring Performance and Flight Morphology 79
6.3.1 Circling Performance .... 79
6.3.2 Cross-Country Soaring ... 81
6.3.3 Wing Shape in Soaring Birds 81
6.4 Bats and Pterosaurs ..... 84
X
Chapter 7. Migration 85
7.1 Introduction . 85
7.2 Orientation and Navigation 85
7.3 Flight Range . . . . . . . . 87
7.4 Cruising Speed and Flight Time 88
7.5 Effect of Wind . . . . . . . . . 90
7.6 Evolution of Soaring Migration 91
7.7 Formation Flight ...... . 93
Chapter 8. Hovering Flight 99
8.1 Introduction . . . . 99
8.2 Kinematics of Hovering 101
8.3 The Rankine-Froude Momentum Theory 103
8.4 Blade-Element Theory ........ . 105
8.4.1 Profile, Parasite and Inertial Power .. . 105
8.4.2 Weis-Fogh's Model for Normal Hovering 106
8.4.3 Norberg's Model for Asymmetrical Hovering 109
8.5 Vortex Theory ...... . 111
8.5.1 Ellington's Hovering Model ... 113
8.5.2 Rayner's Hovering Model . . . . 113
8.6 Animals with Sustained Hovering 115
8.7 Summary and for Ecologists and Others: Recipes for Power
Calculation .............. . 115
8.7.1 Induced Power, Normal Hovering. . . . 116
8.7.2 Induced Power, Asymmetrical Hovering 116
8.7.3 Profile Power, Normal Hovering . . . 116
8.7.4 Profile Power, Asymmetrical Hovering 117
8.7.5 Inertial Power . . . . . . . . . . . . . 117
Chapter 9. Forward Flight 118
9.1 Introduction . . . 118
9.2 Wing Kinematics 118
9.3 Relative Airspeeds and Forces 127
9.3.1 Downstroke Forces 127
9.3.2 Upstroke Forces ....... . 129
9.4 Vorticity Action ....... . 131
9.5 Power Requirements for Horizontal Forward Flight 133
9.5.1 Induced Power 134
9.5.2 Profile Power . 135
9.5.3 Parasite Power 137
9.5.4 Inertial Power . 138
XI
9.5.5 Flapping Flight with Constant Circulation . . . . . . . . . . . 138
9.5.6 A Method of Calculating Forces with Blade-Element Theory. 140
9.5.7 Comparison Between Different Power Models 143
9.6 Take-Off, Climbing and Landing 146
9.6.1 Take-Off and Climbing 147
9.6.2 Landing ..... . 150
9.7 Flight Manoeuvres .. 152
9.7.1 Turning Ability .... 152
9.7.2 Maximum Roll Acceleration and the Initiation of a Tum 152
9.7.3 Prey Catching al).d Landing Manoeuvres 154
9.8 Energy-Saving Types of Flight . 157
9.8.1 Bounding Flight . 157
9.8.2 Undulating Flight . . . . . . . 160
9.8.3 Ground Effect. . . . . . . . . 162
9.9 For Ecologists and Others: Recipes for Power Calculation 164
9.9.1 Induced Power 164
9.9.2 Profile Power . . . . . . . . 164
9.9.3 Parasite Power ....... 165
9.9.4 Power Required for a Climb. 165
9.9.5 Power Required for a Descent 165
9.9.6 Power Required for Bounding Flight 165
9.9.7 Power Required for Undulating Flight 165
Chapter 10. Scaling . . . . . 166
10.1 Introduction..... 166
10.2 Geometric Similarity 167
10.3 Estimated Relationships Between Wing Characteristics and
Body Mass . 168
10.3.1 Wingspan . . 170
10.3.2 Wing Area. . 171
10.3.3 Wing Loading 172
10.3.4 Aspect Ratio 172
10.3.5 Flight Speed, Power and Cost of Transport 174
10.3.6 Flight Muscle Masses . . . . 175
10.3.7 Wingbeat Frequency .... 176
10.4 Upper and Lower Size Limits 177
Chapter 11. Morphological Adaptations for Flight 180
11.1 Introduction .. 180
11.2 Muscle System. 180
XII