Table Of Content

E cological and Environmental Physiology of Birds This page intentionally left blank E cological and Environmental Physiology of Birds J. Eduardo P. W. Bicudo D epartamento de Fisiologia, Instituto de Biociências, Universidade de São Paulo W illiam A. Buttemer S chool of Biological Sciences, University of Wollongong M ark A. Chappell U niversity of California, Riverside J ames T. Pearson D epartment of Physiology, Monash University C laus Bech D epartment of Biology, NTNU, Norway 1 1 Great Clarendon Street, Oxford ox 6dp O xford University Press is a department of the University of Oxford. I t furthers the University’s objective of excellence in research, scholarship, a nd education by publishing worldwide in O xford New York A uckland Cape Town Dar es Salaam Hong Kong Karachi K uala Lumpur Madrid Melbourne Mexico City Nairobi N ew Delhi Shanghai Taipei Toronto W ith offi ces in A rgentina Austria Brazil Chile Czech Republic France Greece G uatemala Hungary Italy Japan Poland Portugal Singapore S outh Korea Switzerland T ailand Turkey Ukraine Vietnam O xford is a registered trade mark of Oxford University Press i n the UK and in certain other countries P ublished in the United States b y Oxford University Press Inc., New York © J. Eduardo P. W. Bicudo, William A. Buttemer, M ark A. Chappell, James T. Pearson, and Claus Bech 2010 T e moral rights of the author have been asserted D atabase right Oxford University Press (maker) F irst published 2010 A ll rights reserved. No part of this publication may be reproduced, s tored in a retrieval system, or transmitted, in any form or by any means, w ithout the prior permission in writing of Oxford University Press, o r as expressly permitted by law, or under terms agreed with the appropriate r eprographics rights organization. Enquiries concerning reproduction o utside the scope of the above should be sent to the Rights Department, O xford University Press, at the address above Y ou must not circulate this book in any other binding or cover a nd you must impose the same condition on any acquirer B ritish Library Cataloguing in Publication Data D ata available L ibrary of Congress Cataloging in Publication Data D ata available T ypeset by SPI Publisher Services, Pondicherry, India P rinted in Great Britain on acid-free paper by t he MPG Books Group in the UK. I SBN 978–0–19–922844–7 (Hbk.) 978–0–19–922845–4 (Pbk.) 1 3 5 7 9 10 8 6 4 2 This page intentionally left blank E cological and Environmental Physiology Series (EEPS) S eries Editor: Warren Burggren, University of North Texas T is authoritative series of concise, aff ordable volumes provides an integrated overview of the ecological and environmental physiology of key taxa including birds, mammals, reptiles, amphibians, insects, crustaceans, mollusks, and fi sh. Each volume provides a state-of-the-art review and synthesis of topics that are relevant to how that specifi c group of organisms have evolved and coped with the environmental characteristics of their habitats. T e series is intended for stu- dents, researchers, consultants, and other professionals in the fi elds of physiology, physiological ecology, ecology, and evolutionary biology. A Series Advisory Board assists in the commissioning of titles and authors, development of volumes, and promotion of the published works. T is Board comprises more than 50 internationally recognized experts in ecological and environmental physiology, providing a combination of both depth and breadth to proposal evaluation and series oversight. T e reader is encouraged to visit the EEPS website for additional information and the latest volumes (h ttp://www.eeps-oxford.com/) . If you have ideas for new titles in this series or just wish to comment on EEPS, please do not hesitate to contact the Series Editor, Warren Burggren (University of North Texas; burg- C ontents A cknowledgements xii F oreword xiii 1 . Introduction — Blueprint of a Bird ( Bauplan/ Body plan) 1 1 .1 What is a Bird? 1 1 .2 Evolution of Birds 2 1 .3 Summary of Passerine Bird Phylogeny 6 1 .4 Evolution of Feathers 8 1 .5 Developmental Biology of Feathers 10 1 .6 Origin and Evolution of Flight 12 1 .7 Energy Requirements for Flight 15 1 .8 Migration 20 M odes of Locomotion 20 M olting 20 M igration Flocks 21 N octurnal Migration 21 E volutionary Aspects 22 A daptations 23 1 .9 Flightlessness 25 1 .10 Overview 27 2 . General Physiological Principles 28 2 .1 Gas Dif usion 28 2 .2 Gas Exchange 29 T he Bird Lung 30 T he Bird Egg 32 2 .3 Temperature and Thermal Exchanges 34 R adiation 35 C onduction 36 C onvection 37 viii Contents E vaporation 38 H eat Balance 39 T hermoneutrality 40 2 .4 Body Size 41 2 .5 Water and Ion Fluxes 43 O smosis 44 B ird Regulation of Water and Ion Fluxes 45 S alt Glands 48 3 . Physiological Bases of Fecundity/Longevity Tradeof s 49 3 .1 Costs of Egg Laying 49 N utrient and Energy Demands 49 S elf-Maintenance/Egg Production Tradeof s 54 F itness Consequences 56 3 .2 Male Mating Costs 59 T estosterone Phenology 59 B ehavioral Ef ects of Testosterone 62 P hysiological Ef ects of Testosterone 64 3 .3 Costs of Parenting 75 C osts of Incubation 76 C osts of Chick Rearing 80 3 .4 Oxidative Stress and Longevity 86 O xidative Stress 86 R eactive O S pecies Production 87 2 A ntioxidant Defenses 92 M embrane Lipids and Peroxidation Resistance 93 4 . Adaptations: Obtaining and Processing Food 99 4 .1 Energy and Nutrition 102 4 .2 Energy and Structure and Function 105 4 .3 Digestive Adaptation 106 C hemical Reactor Theory 107 4 .4 Digestive Ef ciency 110 P henotypic Flexibility 112 4 .5 Modulation of Digestive Enzymes 113 Contents i x 4 .6 Modulation of Absorption Rate 115 4 .7 Modulation of Retention Time 118 G eometric Analysis 120 4 .8 Digestion Rate and Digestive Ef ciency 120 4 .9 Handling and Digestion 123 4 .10 Morphological Adjustments of the Gastrointestinal Tract 127 4 .11 Adjustments during Reproduction 128 4 .12 Adjustments in Migrant Birds 128 4 .13 Plant Secondary Metabolites 131 5 . Adaptations: Living in Specifi c Environments 134 5 .1 Desert and Arid Environments 134 E vaporative Water Loss 136 A rid Environment and Phenotypic Plasticity in Birds 139 H yperthermia 141 C loacal Evaporation 142 E nergy Expenditure and Water Infl ux in the Field 144 C ontrollable Vascular Thermal Radiator 145 5 .2 Low Temperatures 147 M etabolic Heat Production 148 A djustments to Basal Metabolic Rate 148 N on-Shivering Thermogenesis 150 H eat from Digestion 150 H eat Derived from Activity 151 T hermal Conductance 152 H ibernation and Torpor 152 B ehavioral Adaptations 155 A daptations to Low Food Supply 156 C oping with Global Warming? 156 5 .3 High Altitude 157 C onstraints at High Altitude 157 R espiratory Adaptations 160 W ater Loss and Thermoregulation 162 W ind Speeds 165