Evolution

by Ridley, Mark

• ISBN13: 9781405103459
• ISBN10: 1405103450
• Edition: 3rd
• Format: Paperback
• Copyright: 2003-11-07
• Publisher: Wiley-Blackwell

Summary

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 revent 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. Dedicated website - provides an interactive experience of the book, with illustrations downloadable to PowerPoint, and a full supplemental package complementing the book - www.blackwellpublishing.com/ridley. Margin icons - indicate where there is relevant information included in the dedicated website. Two new chapters - one on evolutionary genomics and one on evolution and development bring state-of-the-art information to the coverage of evolutionary study. Two kinds of boxes - one featuring practical applications and the other related information, supply added depth without interrupting the flow of the text. Margin comments - paraphrase and highlight key concepts. Study and review questions - help students review their understanding at the end of each chapter, while new challenge questions prompt students to synthesize the chapter concepts to reinforce the learning at a deeper level.

Author Biography

Mark Ridley works in the Department of Zoology, University of Oxford, UK, and formerly worked at Emory University, Atlanta, and the University of Cambridge, UK. He has carried out research in several areas of evolutionary biology, particularly in sexual selection and the comparative method.

Table of Contents

Preface	xxii
PART 1. INTRODUCTION	1	(92)
The Rise of Evolutionary Biology	3	(18)
Evolution means change in living things by descent with modification	4	(1)
Living things show adaptations	5	(1)
A short history of evolutionary biology	6	(15)
Evolution before Darwin	7	(2)
Charles Darwin	9	(1)
Darwin's reception	10	(4)
The modern synthesis	14	(7)
Summary
Further reading
Study and review questions
Molecular and Mendelian Genetics	21	(22)
Inheritance is caused by DNA molecules, which are physically passed from parent to offspring	22	(1)
DNA structurally encodes information used to build the body's proteins	23	(2)
Information in DNA is decoded by transcription and translation	25	(2)
Large amounts of non-coding DNA exist in some species	27	(1)
Mutational errors may occur during DNA replication	27	(4)
Rates of mutation can be measured	31	(2)
Diploid organisms inherit a double set of genes	33	(1)
Genes are inherited in characteristic Mendelian ratios	34	(3)
Darwin's theory would probably not work if there was a non-Mendelian blending mechanism of heredity	37	(6)
Summary
Further reading
Study and review questions
The Evidence for Evolution	43	(28)
We distinguish three possible theories of the history of life	44	(1)
On a small scale, evolution can be observed in action	45	(2)
Evolution can also be produced experimentally	47	(1)
Interbreeding and phenotypic similarity provide two concepts of species	48	(2)
Ring ``species'' show that the variation within a species can be extensive enough to produce a new species	50	(3)
New, reproductively distinct species can be produced experimentally	53	(1)
Small-scale observations can be extrapolated over the long term	54	(1)
Groups of living things have homologous similarities	55	(6)
Different homologies are correlated, and can be hierarchically classified	61	(3)
Fossil evidence exists for the transformation of species	64	(1)
The order of the main groups in the fossil record suggests they have evolutionary relationships	65	(1)
Summary of the evidence for evolution	66	(1)
Creationism offers no explanation of adaptation	67	(1)
Modern ``scientific creationsim'' is scientifically untenable	67	(4)
Summary
Further reading
Study and review questions
Natural Selection and Variation	71	(22)
In nature, there is a struggle for existence	72	(2)
Natural selection operates if some conditions are met	74	(1)
Natural selection explains both evolution and adaptation	75	(1)
Natural selection can be directional, stablizing, or disruptive	76	(5)
Variation in natural populations is widespread	81	(4)
Organisms in a population vary in reproductive success	85	(2)
New variation is generated by mutation and recombination	87	(1)
Variation created by recombination and mutation is random with respect to the direction of adaptation	88	(5)
Summary
Further reading
Study and review questions
PART 2. EVOLUTIONARY GENETICS	93	(160)
The Theory of Natural Selection	95	(42)
Population genetics is concerned with genotype and gene frequencies	96	(1)
An elementary population genetic model has four main steps	97	(1)
Genotype frequencies in the absence of selection go to the Hardy-Weinberg equilibrium	98	(4)
We can test, by simple observation, whether genotypes in a population are at the Hardy-Weinberg equilibrium	102	(1)
The Hardy-Weinberg theorem is important conceptually, historically, in practical research, and in the workings of theoretical models	103	(1)
The simplest model of selection is for one favored allele at one locus	104	(4)
The model of selection can be applied to the peppered moth	108	(7)
Industrial melanism in months evolved by natural selection	108	(1)
One estimate of the fitnesses is made using the rate of change in game frequencies	109	(2)
A second estimate of the fitnesses is made from the survivorship of the different genotypes in mark-recapture experiments	111	(1)
The selective factor at work is controversial, but bird predation was probably influential	112	(3)
Pesticide resistance in insects is an example of natural selection	115	(3)
Fitnesses are important numbers in evolutionary theory and can be estimated by three main methods	118	(2)
Natural selection operating on a favored allele at a single locus is not meant to be a general model of evolution	120	(1)
A recurrent disadvantageous mutation will evolve to a calculable equilibrial frequency	121	(2)
Heterozygous advantage	123	(4)
Selection can maintain a polymorphism when the heterozygote is fitter than either homozygote	123	(1)
Sickle cell anemia is a polymorphism with heterozygous advantage	124	(3)
The fitness of a genotype may depend on its frequency	127	(2)
Subdivided populations require special population genetic principles	129	(8)
A subdivided set of populations have a higher proportion of homozygotes than an equivalent fused population: this is the Wahlund effect	129	(1)
Migration acts to unify gene frequencies between populations	130	(2)
Convergence of gene frequencies by gene flow is illustrated by the human population of the USA	132	(1)
A balance of selection and migration can maintain genetic differences between subpopulations	132	(5)
Summary
Further reading
Study and review questions
Random Events in Population Genetics	137	(18)
The frequency of alleles can change at random through time in a process called genetic drift	138	(2)
A small funder population may have a non-representative sample of the ancestral population's genes	140	(2)
One gene can be substituted for another by random drift	142	(3)
Hardy-Weinberg ``equilibrium'' assumes the absence of genetic drift	145	(1)
Neutral drift over time produces a march to homozygosity	145	(5)
A calculable amount of polymorphism will exist in a population because of neutral mutation	150	(1)
Population size and effective population size	151	(4)
Summary
Further reading
Study and review questions
Natural Selection and Random Drift in Molecular Evolution	155	(39)
Random drift and natural selection can both hypothetically explain molecular evolution	156	(3)
Rates of molecular evolution and amounts of genetic variation can be measured	159	(5)
Rates of molecular evolution are arguably too constant for a process controlled by natural selection	164	(3)
The molecular clock shows a generation time effect	167	(3)
The nearly neutral theory	170	(5)
The ``purely'' neutral theory faces several empirical problems	170	(1)
The nearly neutral theory of molecular evolution posits a class of nearly neutral mutations	171	(2)
The nearly neutral theory can explain the observed facts better than the purely neutral theory	173	(1)
The nearly neutral theory is conceptually closely related to the original, purely neutral theory	174	(1)
Evolutionary rate and functional constraint	175	(3)
More functionally constrained part of proteins evolve at slower rates	175	(2)
Both natural selection and neutral drift can explain the trend for proteins, but only drift is plausible for DNA	177	(1)
Conclusion and comment: the neutralist paradigm shift	178	(1)
Genomic sequences have led to new ways of studying molecular evolution	179	(11)
DNA sequences provide strong evidence for natural selection on protein structure	180	(1)
A high ratio of non-synonymous to synonymous changes provides evidence of selection	181	(3)
Selection can be detected by comparisons of the dN/dS ratio within and between species	184	(2)
The gene for lysozyme has evolved convergently in cellulosedigesting mammals	186	(1)
Codon usages are biased	187	(2)
Positive and negative selection leave their signatures in DNA sequences	189	(1)
Conclusion: 35 years of research on molecular evolution	190	(4)
Summary
Further reading
Study and review questions
Two-locus and Multilocus Population Genetics	194	(28)
Mimicry in Papilio is controlled by more than one genetic locus	195	(2)
Genotypes at different loci in Papilio memnon are coadapted	197	(1)
Mimicry in Heliconius is controlled by more than one gene, but they are not tightly linked	197	(2)
Two-locus genetics is concerned with haplotype frequencies	199	(1)
Frequencies of haplotypes may or may not be in linkage equilibrium	199	(4)
Human HLA genes are a multilocus gene system	203	(1)
Linkage disequilibrium can exist for several reasons	204	(2)
Two-locus models of natural selection can be built	206	(4)
Hitch-hiking occurs in two-locus selection models	210	(1)
Selective sweeps can provide evidence of selection in DNA sequences	210	(2)
Linkage disequilibrium can be advantageous, neutral, or disadvantageous	212	(2)
Wright invented the influential concept of an adaptive topography	214	(2)
The shifting balance theory of evolution	216	(6)
Summary
Further reading
Study and review questions
Quantitative Genetics	222	(31)
Climatic changes have driven the evolution of beak size in one of Darwin's finches	223	(3)
Quantitative genetics is concerned with characters controlled by large numbers of genes	226	(2)
Variation is first divided into genetic and environmental effects	228	(3)
Variance of a character is divided into genetic and environmental effects	231	(3)
Relatives have similar genotypes, producing the correlation between relatives	234	(1)
Heritability is the proportion of phenotypic variance that is additive	235	(1)
A character's heritability determines its response to artificial selection	236	(4)
strength of selection has been estimated in many studies of natural populations	240	(2)
Relations between genotype and phenotype may be non-linear, producing remarkable responses to selection	242	(3)
Stabilizing selection reduces the genetic variability of a character	245	(1)
Characters in natural populations subject to stabilizing selection show genetic variation	246	(1)
Levels of genetic variation in natural populations are imperfectly understood	247	(2)
Conclusion	249	(4)
Summary
Further reading
Study and review questions
PART 3. ADAPTATION AND NATURAL SELECTION	253	(92)
Adaptive Explanation	255	(37)
Natural selection is the only known explanation for adaptation	256	(3)
Pluralism is appropriate in the study of evolution, not of adaptation	259	(1)
Natural selection can in principle explain all known adaptations	259	(4)
New adaptations evolve in continuous stages from pre-existing adaptations, but the continuity takes various forms	263	(3)
In Darwin's theory, no special process produces evolutionary novelties	263	(1)
The function of an adaptation may change with little change in its from	264	(1)
A new adaptation may evolved by combining unrelated parts	265	(1)
Genetics of adaptation	266	(4)
Fisher proposed a model, and microscope analogy, to explain why the genetic changes in adaptive evolution will be small	266	(2)
An expanded theory is needed when an organism is not near an adaptive peak	268	(1)
The genetics of adaptation is being studied experimentally	268	(2)
Conclusion: the genetics of adaptation	270	(1)
Three main methods are used to study adaptation	270	(2)
Adaptations in nature are not perfect	272	(14)
Adaptations may be imperfect because of time lags	272	(2)
Genetic constraints may cause imperfect adaptation	274	(1)
Developmental constraints may cause adaptive imperfection	275	(6)
Historic constraints may cause adaptive imperfection	281	(3)
An organism's design may be a trade-off between different adaptive needs	284	(1)
Conclusion: constraints on adaptation	284	(2)
How can we recognize adaptations?	286	(6)
The function of an organ should be distinguished from the effects it may have	286	(1)
Adaptations can be defined by engineering design or reproductive fitness	287	(5)
Summary
Further reading
Study and review questions
The Units of Selection	292	(21)
What entities benefit from the adaptations produced by selection?	293	(1)
Natural selection has produced adaptations that benefit various levels of organization	294	(12)
Segregation distortion benefits one gene at the expense of its allele	294	(1)
Selection may sometimes favor some cell lines relative to other cell lines in the same body	295	(1)
Natural selection has produced many adaptations to benefit organisms	296	(2)
Natural selection working on groups of close genetic relatives is called kin selection	298	(3)
Whether group selection ever produces adaptations for the benefit of groups has been controversial, though most biologists now think it is only a weak force in evolution	301	(4)
Which level in the hierarchy of orgnization levels will evolve adaptations is controlled by which level shows heritbility	305	(1)
Another sense of ``unit of selection'' is the entity whose frequency is adjusted directly by natural selection	306	(4)
The two senses of ``unit of selection'' are compatible: one specifies the entity that generally shows phenotypic adaptations, the other the entity whose frequency is generally adjusted by natural selection	310	(3)
Summary
Further reading
Study and review questions
Adaptations in Sexual Reproduction	313	(32)
The existence of sex is an outstanding, unsolved problem in evolutionary biology	314	(6)
Sex has a 50% cost	314	(1)
Sex is unlikely to be explained by genetic constraint	315	(1)
Sex can accelerate the rate of evolution	316	(2)
Is sex maintained by group selection?	318	(2)
There are two main theories in which sex may have a short-term advantage	320	(7)
Sexual reproduction can enable females to reduce the number of delecterious mutations in their offspring	320	(1)
The mutational theory predicts U > 1	321	(2)
Coevolution of parasites and hosts may produce rapid environmental change	323	(4)
Conclusion: it is uncertain how sex is adaptive	327	(1)
The theory of sexual selection explains many differences between males and females	327	(10)
Sexual characters are often apparently deleterious	327	(1)
Sexual selection acts by male competition and female choice	328	(1)
Females may choose to pair with particular males	329	(2)
Females may prefer to pair with handicapped males, because the male's survival indicates his high quality	331	(1)
Female choice in most models of Fisher's and Zahavi's theories is open ended, and this condition can be tested	332	(1)
Fisher's theory requires heritable variation in the male character, and Zahavi's theory requires heritable variation in fitness	333	(2)
Natural selection may work in conflicting ways on males and females	335	(1)
Conclusion: the theory of sex differences is well worked out but incompletely tested	336	(1)
The sex ratio is a well understood adaptation	337	(4)
Natural selection ususlly favors a 50:50 sex ratio	337	(2)
Sex ratios may be biased when either sons or daughters disproportionately act as ``helpers at the nest''	339	(2)
Different adaptations are understood in different levels of detail	341	(4)
Summary
Further reading
Study and review questions
PART 4. EVOLUTION AND DIVERSITY	345	(176)
Species Concepts and Intraspecific Variation	347	(34)
In practice species are recognized and defined by phenetic characters	348	(2)
Several closely related species concepts exist	350	(5)
The biological species concept	351	(2)
The ecological species concept	353	(1)
The phenetic species concept	354	(1)
Isolating barriers	355	(4)
Isolating barriers prevent interbreeding between species	355	(1)
Sperm or pollen competition can produce subtle prezygotic isolation	356	(1)
Closely related African cichlid fish species are prezygotically isolated by their color patterns, but are not postzygotically isolated	357	(2)
Geographic variation within a species can be understood in terms of population genetic and ecological processes	359	(4)
Geographic variation exists in all species and can be caused by adaptation to local conditions	359	(1)
Geographic variation may also be caused by genetic drift	360	(2)
Geographic variation may take the form of a cline	362	(1)
``Popoulation thinking'' and ``typological thinking'' are two ways of thinking about biological diversity	363	(3)
Ecological influences on the from of a species are shown by the phenomenon of character displacement	366	(1)
Some controversial issues exist between the phenetic, biological, and ecological species concepts	367	(7)
The phenetic species concept suffers from serious theoretical defects	368	(1)
Ecological adaptation and gene flow can provide complementary, or in some cases competing, theories of the integrity of species	369	(4)
Both selection and genetic incompatibility provide explanations of reduced hybrid fitness	373	(1)
Taxonomic concepts may be nominalist or realist	374	(3)
The species category	374	(1)
Categories below the species level	375	(1)
Categories above the species level	376	(1)
Conclusion	377	(4)
Summary
Further reading
Study and review questions
Speciation	381	(42)
How can one species split into two reproductively isolated groups of organisms?	382	(1)
A newly evolving species could theoretically have an allopatric, parapatric, or sympatric geographic relation with its ancestor	382	(1)
Reproductive isolation can evolve as a by-product of divergence in allopatric populations	383	(6)
Laboratory experiments illustrate how separately evolving populations of a species tend incidentally to evolve reproductive isolation	384	(2)
Prezygotic isolation evolves because it is genetically correlated with the characters undergoing divergence	386	(1)
Reproductive isolation is often observed when members of geographically distant populations are crossed	387	(2)
Speciation as a by-product of divergence is well documented	389	(1)
The Dobzhansky-Muller theory of postzgotic isolation	389	(10)
The Dobzhansky-Muller theory is a genetic theory of postzygotic isolation, explaining it by interactions among many gene loci	389	(2)
The Dobzhansky-Muller theory is supported by extensive genetic evidence	391	(1)
The Dobzhansky-Muller theory has broad biological plausibility	392	(2)
The Dobzhansky-Muller theory solves a general problem of ``valley crossing'' during speciation	394	(1)
Postzygotic isolation may have ecological as well as genetic causes	395	(1)
Postzygotic isolation usually follows Haldane's rule	396	(3)
An interim conclusion: two solid generalizations about speciation	399	(1)
Reinforcement	399	(6)
Reproductive isolation may be reinforced by natural selection	399	(2)
Preconditions for reinforcement may be short lived	401	(1)
Empirical tests of reinformcement are inconclusive or fail to support the theory	402	(3)
Some plant species have originated by hybridization	405	(3)
Speciation may occur in non-allopatric populations, either parapatrically or sympatrically	408	(1)
Parapatric speciation	409	(2)
Parapatric speciation begins with the evolution of a stepped cline	409	(2)
Evidence for the theory of parapatric speciation is relatively weak	411	(1)
Sympatric speciation	411	(2)
Sympatric speciation is theoretically possible	411	(1)
Phytophagous insects may split sympatrically by host shifts	412	(1)
Phylogenies can be used to test whether speciation has been sympatric or allopatric	413	(1)
The influence of sexual selection in speciation is one current trend in research	413	(2)
Identification of genes that cause reproductive isolation is another current trend in research	415	(2)
Conclusion	417	(6)
Summary
Further reading
Study and review questions
The Reconstruction of Phylogeny	423	(48)
Phylogenies express the ancestral relations between species	424	(1)
Phyloenies are inferred from morphological characters using cladistic techniques	425	(2)
Homologies provide reliable evidence for phylogenetic inference, and homoplasies provide unreliable evidence	427	(3)
Homologies can be distinguished from homoplasies by several criteria	430	(1)
Derived homologies are more reliable indicators of phylogenetic relations than are ancestral homologies	431	(2)
The polarity of character states can be inferred by several techniques	433	(3)
Outgroup comparison	434	(1)
The fossil record	435	(1)
Other methods	436	(1)
Some character conflict may remain after cladistic character analysis is complete	436	(1)
Molecular sequences are becoming increasingly important in phylogenetic inference, and they have distinct properties	437	(2)
Several statistical techniques exist to infer phylogenies from molecular sequences	439	(10)
An unrooted tree is a phylogeny in which the common ancestor is unspecified	439	(1)
One class of molecular phylogenetic techniques uses molecular distances	440	(2)
Molecular evidence may need to be adjusted for the problem of multiple hits	442	(3)
A second class of phylogenetic techniques uses the principle of parsimony	445	(2)
A third class of phylogenetic techniques uses the principle of maximum likelihood	447	(2)
Distance, parsimony, and maximum likelihood methods are all used, but their popularity has changed over time	449	(1)
Molecular phylogenetics in action	449	(2)
Different molecules evolve at different rates and molecular evidene can be tuned to solve particular phylogenetic problems	449	(2)
Molecular phylogenies can now be produced rapidly, and are used in medical research	451	(1)
Several problems have been encountered in molecular phylogenetics	451	(8)
Molecular sequences can be difficult to align	452	(1)
The number of possible trees may be too large for them all to be analyzed	452	(3)
Species in a phylogeny mayhave diverged too little or too much	455	(1)
Different lineages may evolve at different rates	456	(1)
Paralogous genes may be confused with orthologous genes	457	(1)
Conclusion: problems in molecular phylogenetics	458	(1)
Paralogous genes can be used to root unrooted trees	459	(1)
Molecular evidence successfully challenged paleontological evidence in the analysis of human phylogenetic relations	460	(3)
Unrooted trees can be inferred fro other kinds of evidence, such as chromosomal inversions in Hawaiian fruitflies	463	(3)
Conclusion	466	(5)
Summary
Further reading
Study and review questions
Classfication and Evolution	471	(21)
Biologists classify species into a hierarchy of groups	472	(1)
There are phenetic and phylogenetic principles of classification	472	(2)
There are phenetic, cladistic, and evolutionary schools of classification	474	(1)
A method is needed to judge the merit of a school of classification	475	(1)
Phenetic classification uses distance measures and cluster statistics	476	(3)
Phylogenetic classification uses inferred phylogenetic relations	479	(6)
Hennig's cladism classifies species by their phylogenetic branching relations	479	(2)
Cladists distinguish monophyletic, paraphyletic, and polyphyletic groups	481	(2)
A knowledge of phylogeny does not simply tell us the rank levels in Linnaean classification	483	(2)
Evolutionary classification is a synthesis of phenetic and phylogenetic principles	485	(2)
The principle of divergence explains why phylogeny is hierarchical	487	(2)
Conclusion	489	(3)
Summary
Further reading
Study and review questions
Evolutionary Biogeography	492	(29)
Species have defined geographic distributions	493	(3)
Ecological characteristics of a species limit its geographic distribution	496	(1)
Geographic distributions are influenced by dispersal	496	(1)
Geographic distributions are influenced by climate, such as in the ice ages	497	(3)
Local adaptive rediations occur on island archipelagos	500	(3)
Species of large geographic areas tend to be more closely related to other local species than to ecologically similar species elsewhere in the globe	503	(2)
Geographic distributions are influenced by vicariance events, some of which are caused by plate tectonic movements	505	(7)
The Great American Interchange	512	(5)
Conclusion	517	(4)
Summary
Further reading
Study and review questions
PART 5. MACROEVOLUTION	521	(161)
The History of Life	523	(33)
Fossils are remains of organisms from the past and are preserved in sedimentary rocks	524	(1)
Geological time is conventionally divided into a series of eras, periods, and epochs	525	(4)
Successive geological ages were first recognized by characteristic fossil faunas	525	(1)
Geological time is measured in both absolute and relative terms	526	(3)
The history of life: the Precambrian	529	(6)
The origin of life	529	(2)
The origin of cells	531	(2)
The origin of multicellular life	533	(2)
The Cambrian explosion	535	(3)
Evolution of land plants	538	(2)
Vertebrate evolution	540	(5)
Colonization of the land	540	(2)
Mammals evolved from the reptiles in a long series of small changes	542	(3)
Human evolution	545	(5)
Four main classes of change occurred during hominin evolution	545	(2)
Fossil records show something of our ancestors for the past 4 million years	547	(3)
Macroevolution may or may not be an extrapolated from of microevolution	550	(6)
Summary
Further reading
Study and review questions
Evolutionary Genomics	556	(16)
Our expanding knowledge of genome sequences is making it possible to ask, and answer, questions about the evolution of genomes	557	(1)
The human genome documents the history of the human gene set since early life	558	(1)
The history of duplications can be inferred in a genomic sequence	559	(2)
Genome size can shrink by gene loss	561	(2)
Symbiotic mergers, and horizontal gene transfer, between species influence genome evolution	563	(2)
The X/Y sex chromosomes provide an example of evolutionary genomic research at the chromosomal level	565	(2)
Genome sequences can be used to study the history of non-coding DNA	567	(2)
Conclusion	569	(3)
Summary
Further reading
Study and review questions
Evolutionary Developmental Biology	572	(18)
Changes in development, and the genes controlling development, underlie morphological evolution	573	(1)
The theory of recapitulation is a classic idea (largely discredited) about the relation between development and evolution	573	(5)
Humans may have evolved from ancestral apes by changes in regulatory genes	578	(1)
Many genes that regulate development have been identified recently	579	(1)
Modern developmental genetic discoveries have challenged and clarified the meaning of homology	580	(2)
The Hox gene complex has expanded at two points in the evolution of animals	582	(1)
Changes in the embryonic expression of genes are associated with evolutionary changes in morphology	583	(2)
Evolution of genetic switches enables evolutionary innovation, making the system more ``evolvable''	585	(2)
Conclusion	587	(3)
Summary
Further reading
Study and review questions
Rates of Evolution	590	(23)
Rates of evolution can be expressed in ``darwins,'' as illustrated by a study of horse evolution	591	(5)
How do population genetic, and fossil, evolutionary rates compare?	593	(2)
Rates of evolution observed in the short term can explain speciation over longer time periods in Darwin's finches	595	(1)
Why do evolutionary rates vary?	596	(3)
The theory of punctuated equilibrium applies the theory of allopatric speciation to predict the pattern of change in the fossil record	599	(3)
What is the evidence for punctuated equilibrium and for phyletic gradualism?	602	(4)
A satisfactory test requires a complete stratigraphic record and biometrical evidence	602	(1)
Caribbean bryozoans from the Upper Miocene and Lower Pliocene show a punctuated equilibrial pattern of evolution	603	(2)
Ordovician trilobites show gradual evolutionary change	605	(1)
Conclusion	605	(1)
Evolutionary rates can e measured for non-continuous character changes, as illustrated by a study of ``living fossil'' lungfish	606	(3)
Taxonomic data can be used to describe the rate of evolution of higher taxonomic groups	609	(2)
Conclusion	611	(2)
Summary
Further reading
Study and review questions
Coevolution	613	(30)
Coevolution can give rise to coadaptations between species	614	(2)
Coadaptation suggests, but is not conclusive evidence of, coevolution	616	(1)
Insect-plant coevolution	616	(7)
Coevolution between insects and plants may have driven the diversification of both taxa	616	(2)
Two taxa may show mirror-image phylogenies, but coevolution is only one of several explanations for this pattern	618	(2)
Cophylogenies are not found when phytophagous insects undergo host shifts to exploit phylogenetically unrelated but chemically similar plants	620	(2)
Coevolution between plants and insects may explain the grand pattern of diversification in the two taxa	622	(1)
Coevolutionary relations will often be diffuse	623	(1)
Parasite-host coevolution	623	(9)
Evolution of parasitic virulence	625	(5)
Parasites and their hosts may have cophylogenies	630	(2)
Coevolution can proceed in an ``arms race''	632	(5)
Coevolutionary arms races can result in evolutionary escalation	634	(3)
The probability that a species will go extinct is approximately independent of how long it has existed	637	(1)
Antagonistic coevolution can have various forms, including the Red Queen mode	638	(2)
Both biological and physical hypotheses should be tested on macroevolutionary observations	640	(3)
Summary
Further reading
Study and review questions
Extinction and Radiation	643	(39)
The number of species in a taxon increases during phases of adaptive radiation	644	(2)
Causes and consequences of extinctions can be studied in the fossil record	646	(2)
Mass extinctions	648	(7)
The fossil record of extinction rates shows recurrent rounds of mass extinctions	648	(3)
The best studied mass extinction occurred at the Cretaceous-Tertiary boundary	651	(2)
Several factors can contribute to mass extinctions	653	(2)
Distributions of extinction rates may fit a power law	655	(2)
Changes in the quality of the sedimentary record through time are associatied with changes in the observed extinction rate	657	(1)
Species selection	658	(11)
Characters that evolve within taxa may influence extinction and speciation rates, as is illustrated by snails with planktonic and direct development	658	(6)
Differences in the persistence of ecological niches will influence macroevolutionary patterns	664	(1)
When species selection operates, the factors that control macroevolution differ from the factors that control microevolution	665	(1)
From of species selection may change during mass extictions	666	(3)
One higher taxon may replace another, because of chance, enviornmental change, or competitive replacement	669	(5)
Taxonomi patters through time can provide evidence about the cause of replacements	669	(1)
Two bryozoan groups are a possible example of a competitive replacement	670	(1)
Mammals and dinosaurs are a classic example of independent replacement, but recent molecular evidence has complicated the intrpretation	671	(3)
Species diversity may have increased logistically or exponentially since the Cambrian, or it may have increased little at all	674	(3)
Conclusion: biologists and paleontologists have held a range of views about the importance of mass extinctions in the history of life	677	(5)
Summary
Further reading
Study and review questions
Glossary	682	(8)
Answers to Study and Review Questions	690	(9)
Refrences	699	(34)
Index	733