1 Introduction ................................................. 1
The migrant shorebird story .................................. 1
Bodies express ecology ....................................... 2
What is an organism anyway? .................................. 4
Organization of the book ..................................... 5
Scope and readership ......................................... 7
Acknowledgements ............................................. 8
Part I Basics of organismal design
2 Maintaining the balance of heat, water, nutrients, and
energy ...................................................... 15
Dutch dreamcows do not exist ................................ 15
Hot bodies in the cold ...................................... 16
Thermometers do not measure feelings ........................ 20
Balancing water ............................................. 23
Elements of a body .......................................... 26
Birds are not airplanes ..................................... 28
Shorebird insurance strategies .............................. 30
Dying strategically ......................................... 31
Synopsis .................................................... 32
3 Symmorphosis: principle and limitations of economic design .. 33
A well-trained man, a frog, and a hummingbird ............... 33
Economy of design ........................................... 34
Symmorphosis: the principle and the test .................... 36
Safety factors .............................................. 40
Multiple design criteria .................................... 43
One more problem: the climbing of adaptive peaks ............ 44
In addition to oxygen, fires need fuel too .................. 46
Testing symmorphosis in shorebird food-processing systems ... 47
Synopsis .................................................... 49
Part II Adding environment
4 Metabolic ceilings: the ecology of physiological restraint .. 55
Captain Robert Falcon Scott, sled dogs, and limits to
hard work ................................................... 55
The need for a yardstick .................................... 57
Peaks and plateaus: what is true endurance? ................. 60
Regulation of maximal performance: central, peripheral, or
external? ................................................... 61
Temperature and the allometric scaling constant ............. 65
Protecting long-term fitness assets: factorial scopes and
optimal working capacity revisited .......................... 66
The evolution of laziness ................................... 68
Evolutionary wisdom of physiological constraints ............ 69
Beyond Rubner's legacy: why birds can burn their candle at
both ends ................................................... 70
Hard-working shorebirds ..................................... 72
Synopsis .................................................... 75
5 Phenotypic plasticity: matching phenotypesto environmental
demands ..................................................... 79
Adaptive arm-waving, and more ............................... 79
Use it or lose it ........................................... 81
The dynamic gut ............................................. 84
'Classical' phenotypic plasticity: developmental reaction
norms ....................................................... 85
Seasonal phenotype changes in ptarmigan, deer, and
butterflies ................................................. 90
Environmental variability and predictability, and the
kinds of phenotypic adjustments that make sense ............. 93
Degrees of flexibility ...................................... 95
Direct costs and benefits, their trade-offs, and other
layers of constraint ........................................ 96
Phenotypes of fear .......................................... 98
Plasticity: the tinkerer's accomplishment? ................. 101
Phenotypic flexibility in birds ............................ 102
Synopsis ................................................... 106
Part III Adding behaviour
6 Optimal behaviour: currencies and constraints .............. 109
When the going gets tough, the tough get going ............. 109
Loading leatherjackets ..................................... 110
Better lazy than tired ..................................... 114
More haste less speed ...................................... 115
Oystercatchers pressed for time ............................ 117
Informational constraints: getting to know your
environment ................................................ 119
Informational constraints: getting to know your patch ...... 121
Do updating animals really exist? .......................... 122
The psychology of decision-making .......................... 125
Ideal birds sleep together ................................. 128
Synopsis ................................................... 130
7 Optimal foraging: the dynamic choice between diets,
feeding patches, and gut sizes ............................. 131
Eating more by ignoring food ............................... 131
A hard nut to crack ........................................ 133
It takes guts to eat shellfish ............................. 135
Optimal gizzards ........................................... 140
Synopsis ................................................... 144
Part IV Towards a fully integrated view
8 Beyond the physical balance: disease and predation ......... 147
Running with the Red Queen ................................. 147
The responsive nature of'constitutive' innate immunity ..... 149
Body-building to defy death ................................ 151
Coping with danger ......................................... 155
Predicting carrying capacity in the light of fear .......... 158
Synopsis ................................................... 159
9 Population consequences: conservation and management
of flexible phenotypes ..................................... 161
The Holy Grail of population biology ....................... 161
Dredging out bivalve-rich intertidal flats: a case study
on red knots ............................................... 161
Population consequences for the molluscivores .............. 162
Which individual red knots made it through? ................ 165
Migrant flexibility and speed of migration ................. 166
Global change and phenotypic change: plasticity prevails ... 167
How flexible phenotypes cope with advancing springs ........ 169
When organisms can cope no more: limits to phenotypic
change ..................................................... 172
Synopsis ................................................... 173
10 Evolution in five dimensions: phenotypes first ............. 174
Flexible phenotypes and the study of adaptation ............ 174
Separating the environment from the organism ............... 175
and putting them back together: phenotypes first! .......... 176
Genotypes accommodating environmental information? ......... 176
Enter the tarbutniks, and niche construction ............... 179
Evolution in four or five dimensions? ...................... 180
Context, please! An orchestra in need of a theatre ......... 182
Synopsis ................................................... 184
References .................................................... 185
Name Index .................................................... 219
Subject Index ................................................. 222
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