Evolution

Extensively rewritten and reorganized, this new edition of Evolution--featuring a new coauthor: Mark Kirkpatrick (The University of Texas at Austin)--offers additional expertise in evolutionary genetics and genomics, the fastest-developing area of evolutionary biology. Directed toward an undergraduate audience, the text emphasizes the interplay between theory and empirical tests of hypotheses, thus acquainting students with the process of science. It addresses major themes--including the history of evolution, evolutionary processes, adaptation, and evolution as an explanatory framework--at levels of biological organization ranging from genomes to ecological communities.

For Students

Companion Website
The Evolution, Third Edition, Companion Website features review and study tools to help students master the material presented in the textbook. Access to the site is free of charge, and requires no access code. (Instructor registration is required in order for students to access the quizzes.) The site includes the following resources:

* Chapter Outlines and Summaries: Concise overviews of the important topics covered in each chapter.

* Data Analysis Exercises: Expanded for the third edition, these inquiry-based exercises involve students in working with data and analyzing methods and conclusions from published papers.

* Simulation Exercises: Interactive modules that allow students to explore many of the dynamic processes of evolution, and answer questions based on the results they observe.

* Online Quizzes: Quizzes that cover all the major concepts introduced in each chapter. These quizzes are assignable by the instructor.

* Flashcards & Key Terms: Easy-to-use activities that help students learn all the key terminology introduced in each chapter.

* The complete Glossary

For Instructors

Instructor's Resource Library
The Evolution, Third Edition, Instructor's Resource Library includes a variety of resources to help you develop your course and deliver your lectures. The IRL includes the following resources:

* Textbook Figures and Tables: All the figures (including photographs) and tables from the textbook are provided as JPEGs (both high- and low-resolution), reformatted and relabeled for optimal readability when projected.

* PowerPoint Presentations: For each chapter, all of the chapter's figures and tables are provided in a ready-to-use PowerPoint presentation, making it easy to quickly insert figures into your own lecture presentations.

* Answers to the textbook end-of-chapter Problems and Discussion Topics

* Quiz Questions from the Companion Website

* Data Analysis and Simulation Exercises from the Companion Website, with answers

Online Quizzing
A set of online quizzes is available via the Companion Website. These quizzes can be assigned or released for student self-study, at the instructor's discretion. Instructors can also add their own questions to the quizzing system, to create custom quizzes. Results can be viewed online or downloaded for use in gradebook programs. (Instructor registration is required for student access to the quizzes.)

Douglas J. Futuyma is Distinguished Professor in the Department of Ecology and Evolution at the State University of New York at Stony Brook. He received his B.S. from Cornell University and his Ph. D. in Zoology at the University of Michigan with Lawrence Slobodkin. Dr. Futuyma is the author of three previous editions of Evolution, as well as three editions of its predecessor, Evolutionary Biology. He received the 1997 Sewall Wright Award of the American Society of Naturalists and the 2012 Joseph Leidy Award of the Academy of Natural Sciences of Drexel University (Philadelphia). Dr. Futuyma has served as President of the Society for the Study of Evolution, the American Society of Naturalists, and the American Institute of Biological Sciences, and was elected a Fellow of the American Academy of Arts and Sciences in 1996 and the National Academy of Sciences in 2006. He has served as Editor of Evolution and is currently Editor of the Annual Review of Ecology, Evolution, and Systematics. In 2013, he was recognized as Honorary Doctor by the National University of Mongolia. An avid naturalist, his major research interests include evolution of interactions among insects and plants, speciation, and evolution of community structure.

Mark Kirkpatrick is the Painter Centennial Professor of Genetics in the Department of Integrative Biology at the University of Texas at Austin. He received his B.A. in Biology from Harvard in 1978 and his Ph.D. in Zoology from the University of Washington with Monty Slatkin in 1983. Dr. Kirkpatrick has received a Guggenheim Fellowship (1997) and a Poste Rouge Fellowship (France, 1997). He is a Fellow of the American Academy of Arts and Sciences (2008), and of the American Association for the Advancement of Science (2016). Dr. Kirkpatrick received the Sewall Wright Award from the American Society of Naturalists (2014). He has served as Associate Editor of The American Naturalist, Theoretical Population Biology, and Genetics, and on the Editorial Boards of The Annual Review of Ecology, Evolution, and Systematics and Proceedings of the Royal Society of London. Dr. Kirkpatrick's research interests are in evolutionary genetics. He has worked on sexual selection, quantitative genetics, speciation, and species ranges. Current research topics include the evolution of sex determination and chromosome rearrangements.

1. Evolutionary Biology
"Nothing in Biology Makes Sense except in the Light of Evolution"
What Is Evolution? Is It Fact or Theory?
The Evolution of Evolutionary Biology
Before Darwin
Charles Darwin
Darwin's evolutionary theory
Evolutionary biology after Darwin
The evolutionary synthesis
Evolutionary biology since the synthesis
Box 1A. Fundamental Principles of Biological Evolution
How Evolution Is Studied
Philosophical Issues
Ethics, religion, and evolution
Summary

2. The Tree of Life
The Tree of Life, from Darwin to Today
Box 2A. Classification, Taxonomic Practice, and Nomenclature
Phylogenetic Trees
Inferring phylogenies: An introduction
Variations on the Phylogenetic Theme
Branches of a phylogenetic tree sometimes rejoin
Not only organisms have "phylogenies"
Phylogenetic Insights into Evolutionary History
Inferring the history of character evolution
Estimating time of divergence
Patterns of evolution
Box 2B. Evidence for Evolution
Summary

3. Natural Selection and Adaptation
Adaptive Evolution Observed
Natural Selection
The meaning of natural selection
Natural selection and chance
The effective environment depends on the organism
Levels of Selection
Selfish genes and unselfish behaviors
Selection of organisms and groups
Species selection
The Nature of Adaptations
Selection of and selection for
Recognizing adaptations
Imperfections and Constraints
Natural Selection and the Evolution of Diversity
What Not To Expect of Natural Selection
Summary

4. Mutation and Variation
The Machinery of Inheritance
The Inheritance of Variation
Gene mixing by segregation
Gene mixing by recombination
Gene mixing with asexual inheritance
Mutation: The Ultimate Source of Variation
Point mutations
Structural mutations
Rates and Effects of Mutations
Mutation rates
Box 4A. Estimating Mutation Rates
Effects of mutations
Germ line mutations and somatic mutations
Is Mutation Random?
Nongenetic Inheritance
Summary

5. The Genetical Theory of Natural Selection
Natural Selection and Evolution in Real Time
Evolution by Selection and Inheritance
Fitness: The Currency of Selection
Positive Selection: The Spread of Beneficial Mutations
Box 5A. Evolution by Selection on a Single Locus
The rate of adaptation
Chance and adaptation: The probability that a beneficial mutation spreads
Evolutionary Side Effects
Hitchhiking: When one allele goes for a ride with another
When Selection Preserves Variation
Overdominance
Other forms of balancing selection
Selection That Favors the Most Common
Underdominance: When heterozygotes suffer
Positive frequency-dependent selection
The Evolution of a Population's Mean Fitness
The fundamental theorem of natural selection and the adaptive landscape
Deleterious Mutations
A mutation-selection balance
The mutation load
Summary

6. Phenotypic Evolution
Genotypes and Phenotypes
Fitness Functions Describe Selection on Quantitative Traits
Measuring the Strength of Directional Selection
Evolution by Directional Selection
When genes interact: Dominance and epistasis
Adaptation from standing genetic variation versus new mutations
Can adaptation rescue species from extinction?
Artificial Selection
Correlated Traits
Constraints and trade-offs
The causes of genetic correlations
Phenotypic Plasticity
The Genetic Architecture of Quantitative Traits
Quantitative trait loci
The genetics of quantitative traits
Summary

7. Genetic Drift: Evolution at Random
What Is Random Genetic Drift?
The Genealogy of Genes
How Strong Is Genetic Drift?
Populations that change in size
Drift and Genetic Variation within Species
Estimating population size
Genetic Drift and Natural Selection
Crossing an adaptive valley by drift
The fate of beneficial mutations in large populations
The Evolution of Differences among Species
The neutral theory of molecular evolution
Searching the Genes for Signatures of Adaptation
Synonymous versus nonsynonymous differences
The MK test
Divergence among populations
Summary

8. Evolution in Space
Patterns in Space
Gene Flow
How is gene flow measured?
Genetic Divergence between Populations
Gene Flow and Selection
Tension zones
Gene Flow and Drift
Gene flow, local adaptation, and drift
The Evolution of Dispersal
The Evolution of Species' Ranges
Summary

9. Species and Speciation
What Are Species?
Box 9A. Diagnosis of a New Species
Reproductive Isolation
Prezygotic barriers
Postzygotic barriers
How fast does reproductive isolation evolve?
The Causes of Speciation
Box 9B. Speciation in the Lab
The Geography of Speciation
Allopatric speciation
Sympatric speciation
Parapatric speciation
The Genomics of Speciation
Summary

10. All About Sex
What Are Females and Males?
Sexual Selection
Why are males sexually selected?
Sexual selection by male-male competition
Sexual selection by female choice
Sexual selection in flowering plants
Sex Ratios
Why Sex?
Advantages to sex in changing environments
Selective interference favors sex and recombination
Selfing and Outcrossing
Summary

11. How to Be Fit
Life History Traits as Components of Fitness
Costs of reproduction
Fitness in age-structured populations
Senescence
Evolution of the Population Growth Rate and Density
Diverse life histories
Number of offspring
Life histories and mating strategies
Specialists and Generalists
Advantages of specialization
Specialization without trade-offs
Experiments on niche evolution
Summary

12. Cooperation and Conflict
The Costs and Benefits of Interacting
Social Interactions and Cooperation
Cooperation among Unrelated Individuals
Reciprocity
Box 12A. Evolutionarily Stable Strategies
Shared Genes and the Evolution of Altruism
Box 12B. Calculating Relatedness
Box 12C. Altruistic Mating Displays in Turkeys
Spite
Conflict and Cooperation in Close Quarters: The Family
Conflict between mates
Murder in the family
Parent-offspring conflict
Eusocial animals: The ultimate families
Levels of Selection
Selfish DNA
Selfish mitochondria
Group selection
Cooperation and Major Evolutionary Transitions
Summary

13. Interactions among Species
Coevolution and Interactions among Species
The Evolution of Enemies and Victims
Aposematism and mimicry
Plants and herbivores
Parasite-host interactions and infectious disease
Mutualisms
The Evolution of Competitive Interactions
Evolution and Community Structure
Summary

14. The Evolution of Genes and Genomes
The Birth of a Gene
Gene families
The Death of a Gene
Evolution of Protein-Coding Genes
Evolution of coding regions by genetic drift
Evolution of coding regions by positive selection
Evolution of Gene Expression
Gene Structure
Chromosome Evolution
Fissions, fusions, and the evolution of chromosome number
Inversions and the evolution of chromosome structure
Evolution of Genome Size and Content
Genomes large and small
Genetic parasites and transposable elements
Routes to the evolution of the smallest and largest genomes
Summary

15. Evolution and Development
Comparative Development and Evolution
Gene Regulation
Box 15A. Some Methods in Developmental Genetics
Hox genes and the genetic toolkit
Developmental-Genetic Bases of Phenotypic Evolution
Evolution by cis-regulatory mutations
Evolution by trans-regulatory mutations
Overview: The genetics and development of phenotypic evolution
Evolvability and Developmental Pathways
Constraints on Adaptive Evolution
Phenotypic Plasticity and Canalization
Does phenotypic plasticity contribute to evolution?
Summary

16. Phylogeny: The Unity and Diversity of Life
Inferring Phylogenies
Why estimating phylogenies can be hard
Methods for estimating phylogenies
Box 16A. Estimating Trees with Likelihood
How Do We Use Phylogenies?
Dating evolutionary events
Discovering the history of genes and cultures
Reconstructing ancestors
Studying adaptations: The comparative method
Classification
Summary

17. The History of Life
Some Geological Fundamentals
The fossil record
Before Life Began
The Emergence of Life
Precambrian Life
The Cambrian Explosion and the Origins of Animal Diversity
Paleozoic Life
The colonization of land
Paleozoic life on land
The end-Permian mass extinction
Mesozoic Life
The Cenozoic Era
The modern world takes shape
The adaptive radiation of mammals
Pleistocene events
Summary

18. The Geography of Evolution
Biogeographic Evidence for Evolution
Major Patterns of Distribution
Historical factors affecting geographic distributions
Historical Explanations of Geographic Distributions
Vicariance
Dispersal
Phylogeography
Geographic Range Limits: Ecology and Evolution
Geographic Patterns of Diversity
Summary

19. The Evolution of Biological Diversity
Estimating and Modeling Changes in Biological Diversity
Studying diversity in the fossil record
Diversity through the Phanerozoic
Rates of origination and extinction
Mass extinctions
Phylogenetic Studies of Diversity
The shapes of phylogenies
Does Species Diversity Reach Equilibrium?
Summary

20. Macroevolution: Evolution above the Species Level
The Origin of Major New Forms of Life
The origin of mammals
Gradualism and Saltation
The Evolution of Novelty
Incipient and novel features: Permissive conditions and natural selection
Complex characteristics
Homology and the emergence of novel characters
From Microevolution to Macroevolution
Rates of evolution
Gradualism and punctuated equilibria
Speciation and phenotypic evolution
Trends, Predictability, and Progress
Trends: Kinds and causes
Are there major trends in the history of life?
Predictability and contingency in evolution
The question of progress
Summary

21. The Evolutionary Story of Homo sapiens
Where Did We Come From?
Our closest living relatives
How humans differ from other apes
Our ancestry: Hominins through time
The Arrival of Homo sapiens
The human history of hybridization
The diversity of human populations
Brain and Language
Diet and Agriculture: A Revolution in Our World
Box 21A. Domesticated Plants and Animals
Natural Selection, Past and Present
Our genetic loads
Natural selection and evolution in real time
Evolutionary mismatches
The Evolution of Culture
Summary

22. Evolution and Society
Box 22A. Refuting Antievolutionary Arguments
Creationism and Science
Creationism
The nature of science
The Evidence for Evolution
The fossil record
Phylogenetic and comparative studies
Genes and genomes
Biogeography
Failures of the argument from design
Evolution, and its mechanisms, observed
The Uses and Implications of Evolutionary Science
Evolution by natural selection: A broad and flexible concept
Practical applications of evolutionary science
Using organisms' adaptations
Agriculture and natural resources
Conservation
Box 22B. The Current Extinction Crisis
Health and medicine
Evolution and Human Behavior
Variation in cognitive and behavioral traits
Human behavior: Evolution and culture
Understanding nature and humanity
Summary

Appendix: A Statistics Primer
Glossary
Literature Cited
Illustration Credits