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THIRD EDITION
Vertebrate
Life
F Harvey Pough
John B. Heiser
William N. McFarland
-
Cornell University
Technische Hochschule Darmstadt
FACHBEREICH 10 -,. BIOLOGIE
1
- Bibliothek SchnittspahnstraBe 10
D-64287 Darmstadt
Inv.-Nr. M%
•
Macmillan Publishing Company
t
I
New York
Collier Macmillan Publishers
London
Contents
PART ONE
Vertebrate Diversity, Function, and Evolution
1
|
I
The Diversity, Evolution, and Classification
of Vertebrates 3
"^
The Vertebrate Story
3
The Different Kinds of Vertebrates
4
Evolution
12
Variation and Evolution
19
Earth History and Vertebrate Evolution
36
Classification of Vertebrates
38
Traditional and Cladistics Classifications
45
Conclusion
51
Boxes: 1-1 Names and Endings
10
1 - 2 Evolving Views of Evolution
13
1-3 A Special Case of Adaptive Radiation: Species Flocks
1—4 Tempo in Evolution: The Molecular Clock Hypothesis
The Origin of Vertebrates
27
31
55
Some Familiar Facts About Vertebrates
55
The Basic Vertebrate Body Plan
57
A Search for the Relatives of the Vertebrates
57
Environment in Relation to the Origin of Vertebrates
74
Conclusion
76
>
Boxes: 2-1 The Human as a Typical Vertebrate
58
2-2 Multiple Views of the Origin of Vertebrates
70
IX
Vertebrate Organ Systems and Their Evolution
The Unity of Vertebrate Structure
79
Protection, Support, and Movement
91
Energy Acquisition and Support of Metabolism
104
Homeostasis
115
Coordination and Integration
124
Continuity of Life: The Reproductive System
144
/$
*
Homeostasis and Energetics: Water Balance,
Temperature Regulation, and Energy Use 153
The Internal Environment of Vertebrates
153
Exchange of Water and Ions
154
Responses to Temperature
170
Energy Utilization: Patterns Among Vertebrates
Conclusion
191
184
Geology and Ecology During the Origin
of Vertebrates 195
Earth History, Changing Habits, and Vertebrate Evolution
Continental Positions in the Early Paleozoic
195
The Early Habitat of Vertebrates
199
Early Paleozoic Climates
200
-^
Box 5-1 Why the Continents Move
196
PART TWO
Aquatic Vertebrates: Cartilaginous
and Bony Fishes 203
Contents
/"~T;
Earliest Vertebrates
O
The First Evidence of Vertebrates
205
Living Agnathans
221
What Does "Primitive" Mean?
228
Bone and the Early Vertebrates
229
Boxes: 6-1 Reconstructing the First Vertebrates
6—2 Interrelationships of Early Vertebrates
205
208
216
195
79
The Rise of Jawed Vertebrates and the Radiation
of the Chondrichthyes 233
The First Appearance of Jaws and Unique Gnathostome
Characters
233
Placoderms: The Armored Fishes
237
Chondrichthyes: The Cartilaginous Fishes
239
Boxes: 7-1 Food for Sharks
251
7—2 Electroreception by Elasmobranchs
254
Dominating Life in Water: Teleostomes and the
Major Radiation of Fishes 265
Living in Water
265
The Appearance of Teleostomes
289
Living Actinopterygii: Ray-Finned Fishes
305
Living Sarcopterygii: Lobe-Finned Fishes
325
Boxes: 8-1 Mauthner Neurons and the Actinopterygian Brain:
A Quick Getaway
279
8-2 What a Fish's Ears Tell About its Life
319
Geology and Ecology of the Origin
of Tetrapods 333
Continental Geography in the Late Paleozoic
Devonian Climates
333
Terrestrial Habitats of the Devonian
334
333
PART THREE
,
Terrestrial Ectotherms: Amphibians, Turtles,
Crocodilians, and Squamates
338
Origin and Radiation of Tetrapods in the
Late Paleozoic 341
An Interrupted Story
341
Osteolepiform Fishes
341
Evolution of Terrestrial Vertebrates
345
The Radiation and Diversity of Nonamniotic Paleozoic Tetrapods
The Amniotic Egg
363
Early Amniotes
365
Boxes: 10-1 The Bendable Braincase
342
10-2 Jaw Mechanisms of Paleozoic Tetrapods
366
349
Contents
xi
Salamanders, Anurans, and Caecilians
*
Ji
Amphibians
373
Diversity of Life Histories of Amphibians
386
Amphibian Metamorphosis
404
Water Relations of Amphibians
406
Poison Glands and Other Defense Mechanisms
410
Boxes: 11-1 The Subclass Lissamphibia?
374
11 - 2 The Energy Cost of Vocalization by Frogs
396
11-3 "Eye of Newt and Toe of Frog": Toxins in the Skin
of Amphibians
411
Turtles
*•
373
419
Everyone Recognizes a Turtle
419
Phylogenetic Relationships of Turtles
422
Turtle Structure and Function
423
Ecology and Behavior of Turtles
432
Conservation of Turtles
439
Mesozoic Diapsids: Dinosaurs, Birds, Crocodilians,
and Others 445
The Mesozoic Fauna
445
Phylogenetic Relationships Among Diapsids
445
The Archosauromorpha: Dinosaurs and Their Relatives
449
The Saurischian Dinosaurs and the Origin of Birds
460
The Ornithischian Dinosaurs
474
The Ecology and Behavior of Dinosaurs
482
The Lepidosauromorpha: Ichthyosaurs, Lizards, and Others
490
Other Terrestrial Vertebrates of the Late Mesozoic
498
Late Cretaceous Extinctions
498
Box 13-1 Long-Snouted Fish-Eaters
452
Geology and Ecology of Pangaea: Late Paleozoic
and Mesozoic 507
Continental Geography During the Late Paleozoic and Mesozoic
Climatic Prelude to the Mesozoic: Habitats During the
Late Paleozoic
510
Mesozoic Climates and Habitats
511
Terrestrial Plants and Animals of the Cretaceous
512
Mass Extinctions
515
Xii
Contents
507
The Lepidosaurs: Tuatara,{Lizards,
Amphisbaenians, and Snakes
523
The Squamates
523
The Radiation of Squamates
527
Ecology and Behavior of Squamates
539
560
Thermoregulation and the Ecology and Behavior of Squamates
Boxes: 15-1 The Way of a Snake
536
15-2 Caudal Autotomy: Your Tail or Your Life
548
Ectothermy: A Low-Cost Approach to Life
Vertebrates and Their Environments
573
Characteristics of Stressful Environments
574
The Deep Sea
575
^
Ectotherms in the Heat: Deserts
583
Ectotherms in the Cold: Supercooling, Antifreeze,
and Freeze Tolerance
593
The Role of Ectothermal Tetrapods in Terrestrial Ecosystems
Box 16-1 Doubly Labeled Water
584
573
597
PART FOUR
Terrestrial Endotherms: Birds and Mammals
601
Characters of Birds: Specializations
for Flight 603
Birds as Flying Machines
603
Feathers and Flight
604
Aerodynamics of the Avian Wing Compared to Fixed Airfoils
Body Form and Flight
618
The Hind Limbs and Locomotion
630
The Sensory Systems
638
Boxes: 17-1 High-Flying Birds
624
17-2 Not Hearing Straight: Ear Asymetry of Owls
The Ecology and Behavior of Birds
645
The Evolution of Modern Birds
645
Divergence and Convergence in Feeding
650
Social Behavior and Reproduction
662
Migration and Navigation
680
Boxes: 18-1 Giant Predatory Birds
654
18-2 Built-in Babysitters: Nest Helpers
679
608
640
Contents
Xlll
The Synapsida and the Evolution
of Mammals
691
Terrestrial Vertebrates of the Late Paleozoic
691
The Synapsid Skull
691
Pelycosaurs
691
Therapsids and Therosaurs
696
The First Mammals
704
Box 19-1 The Evolution of the Mammalian Middle Ear
Geology and Ecology of the Cenozoic
Continental Geography During the Cenozoic
Periods of Major Glaciatibn
713
Box 20-1 Are Disasters Periodic?
118
Characteristics of Mammals
702
713
713'
725
The Major Lineages of Mammals
725
The Mammalian Integument
730
Mammalian Food and Feeding Specializations
741
Evolution of the Mammalian Nervous and Sensory Systems
758
Mammalian Reproduction
767
Cenozoic Mammals and Vicariance Biogeography
779
Box 21—1 Herbivores, Microbes, and the Ecology of Digestion
744
% {""%
Body Size, Ecology, and Sociality
J\ y^ * of Mammals 795
i
5
Behavior
795
Population Structure and the Distribution of Resources
795
Advantages of Sociality
801
Body Size, Diet, and the Structure of Social Systems
803
Social Systems Among Primates
810
Boxes: 22-1 The Herbivorous Carnivores
798
22-2 Unprofitable Prey?
809
Socia)
Endothermy: A High-Energy Approach
to Life
821
Costs and Benefits
821
Energy Budgets of Vertebrates
821
Endotherms in the Cold: The Arctic
825
Migration to Avoid Stressful Conditions
829
Torpor as a Response to Low Temperatures and Limited Food
XJV
Contents
831
Endotherms in the Heat: Deserts
838
Conclusion
853
jj
Boxes: 23-1 Waking Up Is Hard Work: The Cost of Arousal
23-2 How Hot Is It?
849
Homo sapiens and the Vertebrates
857
The Origin of Humans
857The Human Race and the Future of Vertebrates
Conclusion
895
879
Glossary
834
900
Author Index
I-1
Subject Index 1-7
Contents
XV