Evolution / Edition 3

Evolution / Edition 3

by Mark Ridley

Mark Ridley’s Evolution has become the premier undergraduate text in the study of evolution. Readable and stimulating, yet well balanced and in-depth, this text tells the story of evolution, from the history of the study to the most recent developments in evolutionary theory.

The third edition of this successful textbook features updates and

See more details below


Mark Ridley’s Evolution has become the premier undergraduate text in the study of evolution. Readable and stimulating, yet well balanced and in-depth, this text tells the story of evolution, from the history of the study to the most recent developments in evolutionary theory.

The third edition of this successful textbook features updates and extensive new coverage. The sections on adaptation and diversity have been reorganized for improved clarity and flow, and a completely updated section on the evolution of sex and the inclusion of more plant examples have all helped to shape this new edition. Evolution also features strong, balanced coverage of population genetics, and scores of new applied plant and animal examples make this edition even more accessible and engaging.

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Table of Contents



Part I: Introduction.

1. The Rise Of Evolutionary Biology.

Evolution Means Change In Living Things By Descent WithModification.

Living Things Show Adaptations.

A Short History Of Evolutionary Biology.


Further Reading.

Study And Review Questions.

2. Molecular And Mendelian Genetics.

Inheritance Is Caused By DNA Molecules, Which Are PhysicallyPassed From Parent To Offspring.

DNA Structurally Encodes Information Used To Build TheBody’s Proteins.

Information In DNA Is Decoded By Transcription AndTranslation.

Large Amounts Of Non-Coding DNA Exist In Some Species.

Mutational Errors May Occur During DNA Replication.

Rates Of Mutation Can Be Measured.

Diploid Organisms Inherit A Double Set Of Genes.

Genes Are Inherited In Characteristic Mendelian Ratios.

Darwin’s Theory Would Probably Not Work If There Was ANon-Mendelian Blending Mechanism Of Heredity.


Further Reading.

Study And Review Questions.

3. The Evidence For Evolution.

We Distinguish Three Possible Theories Of The History OfLife.

On A Small Scale, Evolution Can Be Observed In Action.

Evolution Can Also Be Produced Experimentally.

Interbreeding And Phenotypic Similarity Provide Two Concepts OfSpecies.

Ring “Species” Show That The Variation Within ASpecies Can Be Extensive Enough To Produce A New Species.

New, Reproductively Distinct Species Can Be ProducedExperimentally.

Small-Scale Observations Can Be Extrapolated Over The LongTerm.

Groups Of Living Things Have Homologous Similarities.

Different Homologies Are Correlated, And Can Be HierarchicallyClassified.

Fossil Evidence Exists For The Transformation Of Species.

The Order Of The Main Groups In The Fossil Record Suggests TheyHave Evolutionary Relationships.

Summary Of The Evidence For Evolution.

Creationism Offers No Explanation Of Adaptation.

Modern “Scientific Creationism” Is ScientificallyUntenable.


Further Reading.

Study And Review Questions.

4. Natural Selection And Variation.

In Nature, There Is A Struggle For Existence.

Natural Selection Operates If Some Conditions Are Met.

Natural Selection Explains Both Evolution And Adaptation.

Natural Selection Can Be Directional, Stabilizing, OrDisruptive.

Variation In Natural Populations Is Widespread.

Organisms In A Population Vary In Reproductive Success.

New Variation Is Generated By Mutation And Recombination.

New Variation Created By Recombination And Mutation Is RandomWith Respect To The Direction Of Adaptation.


Further Reading.

Study And Review Questions.

Part II: Evolutionary Genetics.

5. The Theory Of Natural Selection.

Population Genetics Is Concerned With Genotype And GeneFrequencies.

An Elementary Population Genetic Model Has Four Main Steps.

Genotype Frequencies In The Absence Of Selection Go To TheHardy–Weinberg Equilibrium.

We Can Test, By Simple Observation, Whether Genotypes In APopulation Are At The Hardy–Weinberg Equilibrium.

The Hardy–Weinberg Theorem Is Important Conceptually,Historically, In Practical Research, And In The Workings OfTheoretical Models.

The Simplest Model Of Selection Is For One Favored Allele At OneLocus.

The Model Of Selection Can Be Applied To The Peppered Moth.

Pesticide Resistance In Insects Is An Example Of NaturalSelection.

Fitnesses Are Important Numbers In Evolutionary Theory And CanBe Estimated By Three Main Methods.

Natural Selection Operating On A Favored Allele At A SingleLocus Is Not Meant To Be A General Model Of Evolution.

A Recurrent Disadvantageous Mutation Will Evolve To A CalculableEquilibrial Frequency.

Heterozygous Advantage.

The Fitness Of A Genotype May Depend On Its Frequency.

Subdivided Populations Require Special Population GeneticPrinciples.


Further Reading.

Study And Review Questions.

6. Random Events In Population Genetics.

The Frequency Of Alleles Can Change At Random Through Time In AProcess Called Genetic Drift.

A Small Founder Population May Have A Non-Representative SampleOf The Ancestral Population’s Genes.

One Gene Can Be Substituted For Another By Random Drift.

Hardy–Weinberg “Equilibrium” Assumes TheAbsence Of Genetic Drift.

Neutral Drift Over Time Produces A March To Homozygosity.

A Calculable Amount Of Polymorphism Will Exist In A PopulationBecause Of Neutral Mutation.

Population Size And Effective Population Size.


Further Reading.

Study And Review Questionss.

7. Natural Selection And Random Drift In MolecularEvolution.

Random Drift And Natural Selection Can Both HypotheticallyExplain Molecular Evolution.

Rates Of Molecular Evolution And Amounts Of Genetic VariationCan Be Measured.

Rates Of Molecular Evolution Are Arguably Too Constant For AProcess Controlled By Natural Selection.

The Molecular Clock Shows A Generation Time Effect.

The Nearly Neutral Theory.

Evolutionary Rate And Functional Constraint.

Conclusion And Comment: The Neutralist Paradigm Shift.

Genomic Sequences Have Led To New Ways Of Studying MolecularEvolution.

Conclusion: 35 Years Of Research On Molecular Evolution.


Further Reading.

Study And Review Questions.

8. Two-Locus And Multilocus Population Genetics.

Mimicry In Papilio Is Controlled By More Than One GeneticLocus.

Genotypes At Different Loci In Papilio Memnon Are Coadapted.

Mimicry In Heliconius Is Controlled By More Than One Gene, ButThey Are Not Tightly Linked.

Two-Locus Genetics Is Concerned With Haplotype Frequencies.

Frequencies Of Haplotypes May Or May Not Be In LinkageEquilibrium.

Human HLA Genes Are A Multilocus Gene System.

Linkage Disequilibrium Can Exist For Several Reasons.

Two-Locus Models Of Natural Selection Can Be Built.

Hitch-Hiking Occurs In Two-Locus Selection Models.

Selective Sweeps Can Provide Evidence Of Selection In DNASequences.

Linkage Disequilibrium Can Be Advantageous, Neutral, OrDisadvantageous.

Wright Invented The Influential Concept Of An AdaptiveTopography.

The Shifting Balance Theory Of Evolution.


Further Reading.

Study And Review Questions.

9. Quantitative Genetics.

Climatic Changes Have Driven The Evolution Of Beak Size In OneOf Darwin’s Finches.

Quantitative Genetics Is Concerned With Characters Controlled ByLarge Numbers Of Genes.

Variation Is First Divided Into Genetic And EnvironmentalEffects.

Variance Of A Character Is Divided Into Genetic AndEnvironmental Effects.

Relatives Have Similar Genotypes, Producing The CorrelationBetween Relatives.

Heritability Is The Proportion Of Phenotypic Variance That IsAdditive.

A Character's Heritability Determines Its Response To ArtificialSelection.

Strength Of Selection Has Been Estimated In Many Studies OfNatural Populations.

Relations Between Genotype And Phenotype May Be Non-Linear,Producing Remarkable Responses To Selection.

Stabilizing Selection Reduces The Genetic Variability Of ACharacter.

Characters In Natural Populations Subject To StabilizingSelection Show Genetic Variation.

Levels Of Genetic Variation In Natural Populations AreImperfectly Understood.



Further Reading.

Study And Review Questions.

Part III: Adaptation And Natural Selection.

10. Adaptive Explanation.

Natural Selection Is The Only Known Explanation ForAdaptation.

Pluralism Is Appropriate In The Study Of Evolution, Not OfAdaptation.

Natural Selection Can In Principle Explain All KnownAdaptations.

New Adaptations Evolve In Continuous Stages From Pre ExistingAdaptations, But The Continuity Takes Various Forms.

Genetics Of Adaptation.

Three Main Methods Are Used To Study Adaptation.

Adaptations In Nature Are Not Perfect.

How Can We Recognize Adaptations?.


Further Reading.

Study And Review Questions.

11. The Units Of Selection.

What Entities Benefit From The Adaptations Produced BySelection?.

Natural Selection Has Produced Adaptations That Benefit VariousLevels Of Organization.

Another Sense Of “Unit Of Selection” Is The EntityWhose Frequency Is Adjusted Directly By Natural Selection.

The Two Senses Of “Unit Of Selection” AreCompatible; One Specifies The Entity That Generally ShowsPhenotypic Adaptations, The Other The Entity Whose Frequency IsGenerally Adjusted By Natural Selection.


Further Reading.

Study And Review Questions.

12. Adaptations In Sexual Reproduction.

The Existence Of Sex Is An Outstanding, Unsolved Problem InEvolutionary Biology.

There Are Two Main Theories In Which Sex May Have A Short-TermAdvantage.

Conclusion: It Is Uncertain How Sex Is Adaptive.

The Theory Of Sexual Selection Explains Many Differences BetweenMales And Females.

The Sex Ratio Is A Well Understood Adaptation.

Different Adaptations Are Understood In Different Levels OfDetail.


Further Reading.

Study And Review Questions.

Part IV: Evolution And Diversity.

13. Species Concepts And Intraspecific Variation.

In Practice Species Are Recognized And Defined By PheneticCharacters.

Several Closely Related Species Concepts Exist.

Isolating Barriers.

Geographic Variation Within A Species Can Be Understood In TermsOf Population Genetic And Ecological Processes.

“Population Thinking” And “TypologicalThinking” Are Two Ways Of Thinking About BiologicalDiversity.

Ecological Influences On The Form Of A Species Are Shown By ThePhenomenon Of Character Displacement.

Some Controversial Issues Exist Between The Phenetic,Biological, And Ecological Species Concepts.

Taxonomic Concepts May Be Nominalist Or Realist.



Further Reading.

Study And Review Questions.

14. Speciation.

How Can One Species Split Into Two Reproductively IsolatedGroups Of Organisms?.

A Newly Evolving Species Could Theoretically Have An Allopatric,Parapatric, Or Sympatric Geographic Relation With Its Ancestor.

Reproductive Isolation Can Evolve As A By-Product Of DivergenceIn Allopatric Populations.

The Dobzhansky–Muller Theory Of Postzygotic Isolation.

An Interim Conclusion: Two Solid Generalizations AboutSpeciation.


Some Plant Species Have Originated By Hybridization.

Speciation May Occur In Non-Allopatric Populations, EitherParapatrically Or Sympatrically.

Parapatric Speciation.

Sympatric Speciation.

The Influence Of Sexual Selection In Speciation Is One CurrentTrend In Research.

Identification Of Genes That Cause Reproductive Isolation IsAnother Current Trend In Research.



Further Reading.

Study And Review Questions.

15. The Reconstruction Of Phylogeny.

Phylogenies Express The Ancestral Relations Between Species.

Phyogenies Are Inferred From Morphological Characters UsingCladistic Techniques.

Homologies Provide Reliable Evidence For Phylogenetic Inference,And Homoplasies Provide Unreliable Evidence.

Homologies Can Be Distinguished From Homoplasies By SeveralCriteria.

Derived Homologies Are More Reliable Indicators Of PhylogeneticRelations Than Are Ancestral Homologies.

The Polarity Of Character States Can Be Inferred By SeveralTechniques.

Some Character Conflict May Remain After Cladistic CharacterAnalysis Is Complete.

Molecular Sequences Are Becoming Increasingly Important InPhylogenetic Inference, And They Have Distinct Properties.

Several Statistical Techniques Exist To Infer Phylogenies FromMolecular Sequences.

Molecular Phylogenetics In Action.

Several Problems Have Been Encountered In MolecularPhylogenetics.

Paralogous Genes Can Be Used To Root Unrooted Trees.

Molecular Evidence Successfully Challenged PaleontologicalEvidence In The Analysis Of Human Phylogenetic Relations.

Unrooted Trees Can Be Inferred From Other Kinds Of Evidence,Such As Chromosomal Inversions In Hawaiian Fruitflies.



Further Reading.

Study And Review Questions.

16. Classification And Evolution.

Biologists Classify Species Into A Hierarchy Of Groups.

There Are Phenetic And Phylogenetic Principles OfClassification.

There Are Phenetic, Cladistic, And Evolutionary Schools OfClassification.

A Method Is Needed To Judge The Merit Of A School OfClassification.

Phenetic Classification Uses Distance Measures And ClusterStatistics.

Phylogenetic Classification Uses Inferred PhylogeneticRelations.

Evolutionary Classification Is A Synthesis Of Phenetic AndPhylogenetic Principles.

The Principle Of Divergence Explains Why Phylogeny IsHierarchical.



Further Reading.

Study And Review Questions.

17. Evolutionary Biogeography.

Species Have Defined Geographic Distributions.

Ecological Characteristics Of A Species Limit Its GeographicDistribution.

Geographic Distributions Are Influenced By Dispersal.

Geographic Distributions Are Influenced By Climate, Such As InThe Ice Ages.

Local Adaptive Radiations Occur On Island Archipelagos.

Species Of Large Geographic Areas Tend To Be More CloselyRelated To Other Local Species Than To Ecologically Similar SpeciesElsewhere In The Globe.

Geographic Distributions Are Influenced By Vicariance Events,Some Of Which Are Caused By Plate Tectonic Movement.

The Great American Interchange.



Further Reading.

Study And Review Questions.

Part V: Macroevolution.

18. The History Of Life.

Fossils Are Remains Of Organisms From The Past And Are PreservedIn Sedimentary Rocks.

Geological Time Is Divided Into A Series Of Eras, Periods, AndEpochs.

The Cambrian Explosion.

Evolution Of Land Plants.

Vertebrate Evolution.

Human Evolution.

Macroevolution May Or May Not Be An Extrapolated Form OfMicroevolution.


Further Reading.

Study And Review Questions.

19. Evolutionary Genomics.

Our Expanding Knowledge Of Genome Sequences Is Making ItPossible To Ask, And Answer, Questions About The Evolution OfGenomes.

The Human Genome Documents The History Of The Human Gene SetSince Early Life.

The History Of Duplications Can Be Inferred In A GenomicSequence.

Genome Size Can Shrink By Gene Loss.

Symbiotic Mergers, And Horizontal Gene Transfer, Between SpeciesInfluence Genome Evolution.

The X/Y Sex Chromosomes Provide An Example Of EvolutionaryGenomic Research At The Chromosomal Level.

Genome Sequences Can Be Used To Study The History Of Non-CodingDNA.



Further Reading.

Study And Review Questions.

20. Evolutionary Developmental Biology.

Changes In Development, And The Genes Controlling Development,Underlie Morphological Evolution.

The Theory Of Recapitulation Is A Classic Idea (LargelyDiscredited) About The Relation Between Development AndEvolution.

Humans May Have Evolved From Ancestral Apes By Changes InRegulatory Genes.

Many Genes That Regulate Development Have Been IdentifiedRecently.

Modern Developmental Genetic Discoveries Have Challenged AndClarified The Meaning On Homology.

The Hox Gene Complex Has Expanded At Two Points In The EvolutionOf Animals.

Changes In The Embryonic Expression Of Genes Are Associated WithEvolutionary Changes In Morphology.

Evolution Of Genetic Switches Enables Evolutionary Innovation,Making The System More “Evolvable”.



Further Reading.

Study And Review Questions.

21. Rates Of Evolution.

Rates Of Evolution Can Be Expressed In “Darwins,” AsIllustrated By A Study Of Horse Evolution.

Why Do Evolutionary Rates Vary?.

The Theory Of Punctuated Equilibrium Applies The Theory OfAllopatric Speciation To Predict The Pattern Of Change In TheFossil Record.

What Is The Evidence For Punctuated Equilibrium And For PhyleticGradualism?.

Evolutionary Rates Can Be Measured For Non-Continuous CharacterChanges, As Illustrated By A Study Of “Living Fossil”Lungfish.

Taxonomic Data Can Be Used To Describe The Rate Of Evolution OfHigher Taxonomic Groups.



Further Reading.

Study And Review Questions.

22. Coevolution.

Coevolution Can Give Rise To Coadaptations Between Species.

Coadaptation Suggests, But Is Not Conclusive Evidence Of,Coevolution.

Insect–Plant Coevolution.

Coevolutionary Relations Will Often Be Diffuse.

Parasite–Host Coevolution.

Coevolution Can Proceed In An “Arms Race”.

The Probability That A Species Will Go Extinct Is ApproximatelyIndependent Of How Long It Has Existed.

Antagonistic Coevolution Can Have Various Forms, Including TheRed Queen Mode.

Both Biological And Physical Hypotheses Should Be Tested OnMacroevolutionary Observations.


Further Reading.

Study And Review Questions.

23. Extinction And Radiation.

The Number Of Species In A Taxon Increases Durings Phases OfAdaptive Radiation.

Causes And Consequences Of Extinctions Can Be Studied In TheFossil Record.

Mass Extinctions.

Distributions Of Extinction Rates May Fit A Power Law.

Changes In The Quality Of The Sedimentary Record Through TimeAre Associated With Changes In The Observed Extinction Rate.

Species Selection.

One Higher Taxon May Replace Another, Because Of Chance,Environmental Change, Or Competitive Replacement.

Species Diversity May Have Increased Logistically OrExponentially Since The Cambrian, Or It May Have Increased LittleAt All.

Conclusion: Biologists And Paleontologists Have Held A Range OfViews About The Importance Of Mass Extinctions In The History OfLife.


Further Reading.

Study And Review Questions.


Answers To Study And Review Questions.



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