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The Princeton Guide to Evolution Digital Instant
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ISBN(s): 9781400848065, 1400848067
Edition: Course Book
File Details: PDF, 15.58 MB
Year: 2013
Language: english
The Princeton Guide to
Evolution
princeton university press
Princeton & Oxford
The Princeton Guide to
Evolution
editor in chief Jonathan B. Losos
Harvard University
editors David A. Baum
University of Wisconsin, Madison
Douglas J. Futuyma
Stony Brook University
Hopi E. Hoekstra
Harvard University
Richard E. Lenski
Michigan State University
Allen J. Moore
University of Georgia
Catherine L. Peichel
Fred Hutchinson Cancer 3esearch
Center
Dolph Schluter
University of British Columbia
Michael J. Whitlock
University of British Columbia
advisors Michael J. Donoghue
Simon A. Levin
Trudy F. C. Mackay
Loren Rieseberg
Joseph Travis
Gregory A. Wray
Copyright # 2014 by Princeton University Press
Published by Princeton University Press, 41 William Street,
Princeton, New Jersey 08540
In the United Kingdom: Princeton University Press, 6 Oxford Street,
Woodstock, Oxfordshire OX20 1TW
press.princeton.edu
All Rights Reserved
Library of Congress Cataloging-in-Publication Data
The Princeton guide to evolution / Jonathan B. Losos, Harvard
University, editor in chief ; David A. Baum, University of Wisconsin,
Madison, Douglas J. Futuyma, Stony Brook University, Hopi E.
Hoekstra, Harvard University, Richard E. Lenski, Michigan State
University, Allen J. Moore, University of Georgia, Catherine L.
Peichel, Hutchinson Cancer Institute, Seattle, Dolph Schluter,
University of British Columbia, Michael C. Whitlock, University
of British Columbia, editors.
pages cm
Includes bibliographical references and index.
ISBN 978-0-691-14977-6 (hardcover)
1. Evolution (Biology) I. Losos, Jonathan B.
QH367.P85 2014
576.8—dc23
2013022360
British Library Cataloging-in-Publication Data is available
This book has been composed in Sabon and Din
Printed on acid-free paper.
Printed in the United States of America
10 9 8 7 6 5 4 3 2 1
Contents
Preface vii
Contributors ix
Section I Introduction 1
I.1 What Is Evolution? 3
I.2 The History of Evolutionary Thought 10
I.3 The Evidence for Evolution 28
I.4 From DNA to Phenotypes 40
Section II Phylogenetics and the
History of Life 47
II.1 Interpretation of Phylogenetic Trees 51
II.2 Phylogenetic Inference 60
II.3 Molecular Clock Dating 67
II.4 Historical Biogeography 75
II.5 Phylogeography 82
II.6 Concepts in Character Macroevolution:
Adaptation, Homology, and Evolvability 89
II.7 Using Phylogenies to Study Phenotypic
Evolution: Comparative Methods
and Tests of Adaptation 100
II.8 Taxonomy in a Phylogenetic Framework 106
II.9 The Fossil Record 112
II.10 The Origin of Life 120
II.11 Evolution in the Prokaryotic Grade 127
II.12 Origin and Diversification of Eukaryotes 136
II.13 Major Events in the Evolution
of Land Plants 143
II.14 Major Events in the Evolution of Fungi 152
II.15 Origin and Early Evolution of Animals 159
II.16 Major Events in the Evolution of
Arthropods 167
II.17 Major Features of Tetrapod Evolution 174
II.18 Human Evolution 183
Section III Natural Selection
and Adaptation 189
III.1 Natural Selection, Adaptation, and Fitness:
Overview 193
III.2 Units and Levels of Selection 200
III.3 Theory of Selection in Populations 206
III.4 Kin Selection and Inclusive Fitness 215
III.5 Phenotypic Selection on Quantitative
Traits 221
III.6 Responses to Selection: Experimental
Populations 230
III.7 Responses to Selection:
Natural Populations 238
III.8 Evolutionary Limits and Constraints 247
III.9 Evolution of Modifier Genes and
Biological Systems 253
III.10 Evolution of Reaction Norms 261
III.11 Evolution of Life Histories 268
III.12 Evolution of Form and Function 276
III.13 Biochemical and Physiological Adaptations 282
III.14 Evolution of the Ecological Niche 288
III.15 Adaptation to the Biotic Environment 298
Section IV Evolutionary Processes 305
IV.1 Genetic Drift 307
IV.2 Mutation 315
IV.3 Geographic Variation, Population
Structure, and Migration 321
IV.4 Recombination and Sex 328
IV.5 Genetic Load 334
IV.6 Inbreeding 340
IV.7 Selfish Genetic Elements and Genetic
Conflict 347
IV.8 Evolution of Mating Systems: Outcrossing
versus Selfing 356
Section V Genes, Genomes, Phenotypes 363
V.1 Molecular Evolution 367
V.2 Genome Evolution 374
V.3 Comparative Genomics 380
V.4 Evolution of Sex Chromosomes 387
V.5 Gene Duplication 397
V.6 Evolution of New Genes 406
V.7 Evolution of Gene Expression 413
V.8 Epigenetics 420
V.9 Evolution of Molecular Networks 428
V.10 Evolution and Development: Organisms 436
V.11 Evolution and Development: Molecules 444
V.12 Genetics of Phenotypic Evolution 452
V.13 Dissection of Complex Trait Evolution 458
V.14 Searching for Adaptation in the Genome 466
V.15 Ancient DNA 475
Section VI Speciation and Macroevolution 483
VI.1 Species and Speciation 489
VI.2 Speciation Patterns 496
VI.3 Geography, Range Evolution, and
Speciation 504
VI.4 Speciation and Natural Selection 512
VI.5 Speciation and Sexual Selection 520
VI.6 Gene Flow, Hybridization, and Speciation 529
VI.7 Coevolution and Speciation 535
VI.8 Genetics of Speciation 543
VI.9 Speciation and Genome Evolution 549
VI.10 Adaptive Radiation 559
VI.11 Macroevolutionary Rates 567
VI.12 Macroevolutionary Trends 573
VI.13 Causes and Consequences of Extinction 579
V1.14 Species Selection 586
VI.15 Key Evolutionary Innovations 592
VI.16 Evolution of Communities 599
Section VII Evolution of Behavior, Society,
and Humans 605
VII.1 Genes, Brains, and Behavior 609
VII.2 Evolution of Hormones and Behavior 616
VII.3 Game Theory and Behavior 624
VII.4 Sexual Selection and Its Impact on Mating
Systems 632
VII.5 Sexual Selection: Male-Male Competition 641
VII.6 Sexual Selection: Mate Choice 647
VII.7 Evolution of Communication 655
VII.8 Evolution of Parental Care 663
VII.9 Cooperation and Conflict: Microbes
to Humans 671
VII.10 Cooperative Breeding 677
VII.11 Human Behavioral Ecology 683
VII.12 Evolutionary Psychology 690
VII.13 Evolution of Eusociality 697
VII.14 Cognition: Phylogeny, Adaptation,
and By-Products 703
VII.15 Evolution of Apparently Nonadaptive
Behavior 710
VII.16 Aging and Menopause 718
Section VIII Evolution and Modern Society 727
VIII.1 Evolutionary Medicine 733
VIII.2 Evolution of Parasite Virulence 741
VIII.3 Evolution of Antibiotic Resistance 747
VIII.4 Evolution and Microbial Forensics 754
VIII.5 Domestication and the Evolution of
Agriculture 760
VIII.6 Evolution and Conservation 766
VIII.7 Directed Evolution 774
VIII.8 Evolution and Computing 780
VIII.9 Linguistics and the Evolution
of Human Language 786
VIII.10 Cultural Evolution 795
VIII.11 Evolution and Notions of Human Race 801
VIII.12 The Future of Human Evolution 809
VIII.13 Evolution and Religion 817
VIII.14 Creationism and Intelligent Design 825
VIII.15 Evolution and the Media 832
Index 837
vi Contents
Preface
For more than 150 years, since the publication of On the
Origin of Species, biologists have focused on under-
standing the evolutionary chronicle of diversification and
extinction, and the underlying evolutionary processes
that have produced it. Although progress in evolutionary
biology has been steady since Darwin’s time, develop-
ments in the last 20 years have ushered in a golden era of
evolutionary study in which biologists are on the brink of
answeringmanyofthefundamentalquestionsinthefield.
These advances have come from a confluence of tech-
nological and conceptual innovations. In the laboratory,
the rapid and inexpensive sequencing of large amounts
of DNA is producing a wealth of data on the genomes
of many species; comparisons of these genomes are al-
lowing scientists to pinpoint the specific genetic changes
that have occurred over the course of evolution. In par-
allel, spectacular fossil discoveries have filled many of
the most critical gaps in our documentation of the evo-
lutionary pageant, detailing how whales evolved from
land-living animals, snakes from their four-legged lizard
forebears, and humans from our primate ancestors. In
addition, providing the data that Darwin could only
imagine, field biologists are now tracking populations,
directly documenting natural selection as it occurs, and
monitoring the resulting evolutionary changes that oc-
cur from one generation to the next.
At the same time, evolutionary biology is making an
impactthroughouthumansociety.Manycurrentissues—
such as the rise of new diseases, the increasedresistance of
pests and microorganisms to efforts to control them, and
the effect of changing environmental conditions on nat-
ural populations—revolve around aspects of natural se-
lection and evolutionary change. Many disparate areas of
modern life—medicine, the legal system, computing—
increasingly employ evolutionary thinking and use meth-
odsdevelopedinevolutionarybiology.Paradoxically,even
as our understanding of evolution and its importance to
society has never been greater, substantial proportions of
thepopulationinanumberofcountries—mostnotablythe
UnitedStatesandTurkey—disputethescientificfindingsof
evolutionary biologists and resist the teaching of evolution
in schools.
This volume follows on the success of The Princeton
Guide to Ecology, edited by Simon Levin. Published in
2009, the ecology guide has proven valuable to a wide
range of readers, from professional ecologists and
graduate students to land planners, economists, and
social scientists. With this model in mind, we set out to
produce a guide that would be accessible and useful to
students and scientists in evolutionary biology and re-
lated disciplines, as well as to anyone with a serious in-
terest in evolution. What makes this volume stand out is
the breadth and depth of our 107 chapters, each written
by authorities in their respective field. In addition, the
articles balance accessibility with depth of analysis,
making the Guide a valuable reference for a broad au-
dience. Certainly, some articles are more technical than
others, but readers can easily select chapters appropriate
for their interests and expertise.
The Guide is divided into eight sections. The intro-
ductory section includes four chapters covering the basics
of evolution: what it is, the history of its study, the evi-
dence for its occurrence, and a basic primer of genetic and
phenotypic variation. The following seven sections cover
the major areas of evolutionary biology, each beginning
with a synoptic overview by the section editor. Section II:
PhylogeneticsandtheHistoryofLife,coversthehistoryof
life andhowitisstudied.Itincludeschaptersontheevolu-
tion of each of the major forms of life, as well as on the
studyoflife’shistorythroughtheexaminationofthefossil
record and the construction of phylogenetic trees that
detail the relationships among species and higher taxa.
Section III: Selection and Adaptation, moves to evolu-
tionary processes, focusing on natural selection, the pre-
sumed primary driver of evolutionary change. Section IV:
Evolutionary Processes, covers gene flow, genetic drift,
andnonrandommating.SectionV:Genes,Genomes,Phe-
notypes, examines the link between genes and phenotypes
and how they evolve, focusing on the rapid growth of
knowledge and continuing research in genetics and de-
velopmental biology and the relationships of these fields
to evolutionary biology. Section VI: Speciation and Mac-
roevolution, moves the focus to the species level and
above,emphasizingtheoriginofspecies—thatis,speciation
—andevolutionarychangethatdriveslarge-scalechanges
in the history of life through time, such as the rise of
particular taxa and the extinction of others. Section VII:
Evolution of Behavior, Society, and Humans, focuses on
behavioral and social interactions that occur within spe-
cies, including competition for mating success (referredto
as sexual selection) and the evolution of traits such as
parental care, communication, and altruism. Although
chapters in this section are broad in taxonomic scope,
many have particular relevance to human biology. Fi-
nally, Section VIII: Evolution and Modern Society, ad-
dresses how evolutionary biology directly affects the
health and welfare of humans today.
This volume could not have been possible without the
efforts of the editors and authors, whose work was instru-
mental to such a wide-ranging and authoritative work.
Developmentoftheguidealsobenefitedimmenselyfromthe
wisdom of our advisors, Michael Donoghue, Simon Levin,
Trudy Mackay, Loren Rieseberg, Joseph Travis, and Greg
Wray.Inaddition,theeditorialstaffatPrincetonUniversity
Press was indispensable. The entire project was skillfully
overseenbyexecutiveeditorAnneSavarese,andday-to-day
management moved smoothly and efficiently under the
watchful eyes of editorial assistants Diana Goovaerts and
Sarah David, and production editor Karen Carter.
Wemournthelossandgratefullyacknowledgethecon-
tribution of our distinguished colleague Farish Jenkins,
who died on November 11, 2012.
Jonathan B. Losos
viii Preface
Contributors
Aneil F. Agrawal, Department of Ecology and Evolutionary
Biology, University of Toronto
IV.5 GENETIC LOAD
Michael E. Alfaro, Department of Ecology and Evolutionary
Biology, University of California, Los Angeles
VI.15 KEY EVOLUTIONARY INNOVATIONS
Garland E. Allen, Department of Biology, Washington
University in St. Louis
I.2 THE HISTORY OF EVOLUTIONARY THOUGHT
Dan I. Andersson, Department of Medical Biochemistry and
Microbiology, Uppsala University
VIII.3 EVOLUTION OF ANTIBIOTIC RESISTANCE
Michael J. Angilletta Jr., School of Life Sciences, Arizona
State University
III.13 BIOCHEMICAL AND PHYSIOLOGICAL ADAPTATIONS
Charles F. Aquadro, Department of Molecular Biology and
Genetics, Cornell University
V.1 MOLECULAR EVOLUTION
Jonathan W. Atwell, Department of Biology, Indiana
University
VII.2 EVOLUTION OF HORMONES AND BEHAVIOR
Francisco J. Ayala, Department of Ecology and Evolutionary
Biology, University of California, Irvine
VIII.13 EVOLUTION AND RELIGION
Doris Bachtrog, Department of Integrative Biology,
University of California, Berkeley
V.4 EVOLUTION OF SEX CHROMOSOMES
Charles F. Baer, Department of Biology, University of Florida
IV.2 MUTATION
Nathan W. Bailey, School of Biology, University of St.
Andrews
VII.15 EVOLUTION OF APPARENTLY NONADAPTIVE
BEHAVIOR
Timothy G. Barraclough, Division of Ecology and Evolution,
Imperial College London
VI.2 SPECIATION PATTERNS
Spencer C. H. Barrett, Department of Ecology and
Evolutionary Biology, University of Toronto
IV.8 EVOLUTION OF MATING SYSTEMS: OUTCROSSING
VERSUS SELFING
N. H. Barton, Institute of Science and Technology Austria
IV.4 RECOMBINATION AND SEX
David A. Baum, Department of Botany, University of
Wisconsin, Madison
II PHYLOGENETICS AND THE HISTORY OF LIFE
Graham Bell, Department of Biology, McGill University
III.6 RESPONSES TO SELECTION: EXPERIMENTAL
POPULATIONS
Yehuda Ben-Shahar, Department of Biology, Washington
University in St. Louis
VII.1 GENES, BRAINS, AND BEHAVIOR
Michael J. Benton, School of Earth Sciences, University of
Bristol
VI.13 CAUSES AND CONSEQUENCES OF EXTINCTION
Janette W. Boughman, Department of Zoology, Michigan
State University
VI.5 SPECIATION AND SEXUAL SELECTION
Paul M. Brakefield, Department of Zoology, University of
Cambridge
V.10 EVOLUTION AND DEVELOPMENT: ORGANISMS
Edmund D. Brodie III, Department of Biology, University of
Virginia
III.5 PHENOTYPIC SELECTION ON QUANTITATIVE
TRAITS
C. Alex Buerkle, Department of Botany and Program in
Ecology, University of Wyoming
VI.6 GENE FLOW, HYBRIDIZATION, AND
SPECIATION
Michael A. Cant, Biosciences, University of Exeter
VII.10 COOPERATIVE BREEDING
Paulyn Cartwright, Department of Ecology and Evolutionary
Biology, University of Kansas
II.15 ORIGIN AND EARLY EVOLUTION OF ANIMALS
Amy Cavanaugh, Department of Biological Sciences,
University of Wisconsin, Rock County
VIII.5 DOMESTICATION AND THE EVOLUTION OF
AGRICULTURE
Michel Chapuisat, Department of Ecology and Evolution,
University of Lausanne
VII.13 EVOLUTION OF EUSOCIALITY
Deborah Charlesworth, School of Biological Sciences,
University of Edinburgh
IV.6 INBREEDING
Julia Clarke, Jackson School of Geosciences, University of
Texas at Austin
II.8 TAXONOMY IN A PHYLOGENETIC FRAMEWORK
Peter R. Crane, School of Forestry and Environmental
Studies, Yale University
II.13 MAJOR EVENTS IN THE EVOLUTION OF LAND
PLANTS
Cameron R. Currie, Department of Bacteriology, University
of Wisconsin, Madison
VIII.5 DOMESTICATION AND THE EVOLUTION OF
AGRICULTURE
David Deamer, Department of Biomolecular Engineering,
University of California, Santa Cruz
II.10 THE ORIGIN OF LIFE
Michael J. Donoghue, Department of Ecology and
Evolutionary Biology, Yale University
II.4 HISTORICAL BIOGEOGRAPHY
Dieter Ebert, Zoological Institute,
Universität Basel
VIII.2 EVOLUTION OF PARASITE VIRULENCE
Scott P. Egan, Department of Biological Sciences, University
of Notre Dame
VI.9 SPECIATION AND GENOME EVOLUTION
Andrew D. Ellington, Department of Chemistry and
Biochemistry, University of Texas at Austin
VIII.7 DIRECTED EVOLUTION
Jeffrey Feder, Department of Biological Sciences, University
of Notre Dame
VI.9 SPECIATION AND GENOME EVOLUTION
Lila Fishman, Division of Biological Sciences, University of
Montana
IV.7 SELFISH GENETIC ELEMENTS AND GENETIC
CONFLICT
Douglas J. Futuyma, Department of Ecology and Evolution,
Stony Brook University
III NATURAL SELECTION AND ADAPTATION
Dana H. Geary, Department of Geoscience, University of
Wisconsin, Madison
II.9 THE FOSSIL RECORD
J. Peter Gogarten, Department of Molecular and Cell
Biology, University of Connecticut
II.11 EVOLUTION IN THE PROKARYOTIC GRADE
Emma E. Goldberg, Biological Sciences, University of
Illinois, Chicago
VI.14 SPECIES SELECTION
Peter R. Grant, Department of Ecology and Evolutionary
Biology, Princeton University
VI.10 ADAPTIVE RADIATION
Michael D. Greenfield, Institut de Recherche sur la Biologie
de l’Insecte, Université de Tours
VII.7 EVOLUTION OF COMMUNICATION
Elizabeth Hannon, Department of History and Philosophy of
Science, University of Cambridge
VIII.10 CULTURAL EVOLUTION
Sara J. Hanson, Department of Biology and Program in
Genetics, University of Iowa
V.2 GENOME EVOLUTION
Luke J. Harmon, Department of Biological Sciences,
University of Idaho
VI.11 MACROEVOLUTIONARY RATES
Richard G. Harrison, Department of Ecology and
Evolutionary Biology, Cornell University
VI.1 SPECIES AND SPECIATION
Marc D. Hauser, Independent Scholar
VII.14 COGNITION: PHYLOGENY, ADAPTATION, AND BY-
PRODUCTS
John Hawks, Department of Anthropology, University of
Wisconsin, Madison
II.18 HUMAN EVOLUTION
Philip Hedrick, School of Life Sciences, Arizona State
University
IV.1 GENETIC DRIFT
Noel A. Heim, Department of Geology and Evolutionary
Science, Stanford University
II.9 THE FOSSIL RECORD
Michael E. Hellberg, Department of Biological Sciences,
Louisiana State University
II.5 PHYLOGEOGRAPHY
David S. Hibbett, Department of Biology, Clark University
II.14 MAJOR EVENTS IN THE EVOLUTION OF FUNGI
Hopi E. Hoekstra, Department of Organismic and
Evolutionary Biology, Harvard University
V GENES, GENOMES, PHENOTYPES
Ary Hoffmann, Department of Genetics and Zoology,
University of Melbourne
III.8 LIMITS AND CONSTRAINTS
Mark Holder, Department of Ecology and Evolutionary
Biology, University of Kansas
II.2 PHYLOGENETIC INFERENCE
Kent E. Holsinger, Department of Ecology and Evolutionary
Biology, University of Connecticut
III.3 THEORY OF SELECTION IN POPULATIONS
Robert D. Holt, Department of Ecology, University of Florida
III.14 EVOLUTION OF THE ECOLOGICAL NICHE
Robin Hopkins, Department of Integrative Biology,
University of Texas at Austin
VI.4 SPECIATION AND NATURAL SELECTION
Gene Hunt, Department of Paleobiology Smithsonian
Institution, National Museum of Natural History
VI.12 MACROEVOLUTIONARY TRENDS
John Jaenike, Department of Biology, University of
Rochester
IV.7 SELFISH GENETIC ELEMENTS AND GENETIC CONFLICT
Farish A. Jenkins Jr., Late Professor of Biology, Harvard
University
II.17 MAJOR FEATURES OF TETRAPOD EVOLUTION
Michael D. Jennions, Research School of Biology, Australian
National University
VII.6 SEXUAL SELECTION: MATE CHOICE
Laura A. Katz, Department of Biological Sciences, Smith
College
II.12 ORIGIN AND DIVERSIFICATION OF EUKARYOTES
x Contributors
Paul Keim, Department of Biology, Northern Arizona
University
VIII.4 EVOLUTION AND MICROBIAL FORENSICS
Laurent Keller, Department of Ecology and Evolution,
University of Lausanne
VII.13 EVOLUTION OF EUSOCIALITY
Ellen D. Ketterson, Department of Biology, Indiana
University
VII.2 EVOLUTION OF HORMONES AND BEHAVIOR
Joel G. Kingsolver, Department of Biology, University of
North Carolina, Chapel Hill
III.7 RESPONSES TO SELECTION: NATURAL
POPULATIONS
Hanna Kokko, Research School of Biology, Australian
National University
VII.6 SEXUAL SELECTION: MATE CHOICE
Mathias Kölliker, Department of Environmental Sciences,
University of Basel
VII.8 EVOLUTION OF PARENTAL CARE
Allan Larson, Department of Biology, Washington
University in St. Louis
II.6 CONCEPTS IN CHARACTER MACROEVOLUTION:
ADAPTATION, HOMOLOGY, AND EVOLVABILITY
Richard E. Lenski, Departments of Microbiology &
Molecular Genetics, Zoology, and Crop and Soil Sciences,
Michigan State University
VIII EVOLUTION AND MODERN SOCIETY
Andrew B. Leslie, School of Forestry and Environmental
Studies, Yale University
II.13 MAJOR EVENTS IN THE EVOLUTION OF LAND
PLANTS
Tim Lewens, Department of History and Philosophy of
Science, University of Cambridge
VIII.10 CULTURAL EVOLUTION
John M. Logsdon Jr., Department of Biology and Program in
Genetics, University of Iowa
V.2 GENOME EVOLUTION
Manyuan Long, Department of Ecology and Evolution,
University of Chicago
V.6 EVOLUTION OF NEW GENES
Jonathan B. Losos, Department of Organismic and
Evolutionary Biology, Harvard University
I.1 WHAT IS EVOLUTION?
David B. Lowry, Department of Integrative Biology,
University of Texas at Austin
VI.4 SPECIATION AND NATURAL SELECTION
Virpi Lummaa, Department of Animal and Plant Sciences,
University of Sheffield
VII.11 HUMAN BEHAVIORAL ECOLOGY
Florian Maderspacher, Current Biology, Elsevier, Inc.
V.8 EPIGENETICS
Gregory C. Mayer, Department of Biological Sciences,
University of WisconsinParkside
I.3 THE EVIDENCE FOR EVOLUTION
Joel W. McGlothlin, Department of Biological Sciences,
Virginia Polytechnic Institute and State University
VII.2 EVOLUTION OF HORMONES AND BEHAVIOR
Daniel McNabney, Department of Biology, University of
Rochester
VI.8 GENETICS OF SPECIATION
John M. McNamara, School of Mathematics,
University of Bristol
VII.3 GAME THEORY AND BEHAVIOR
Mark A. McPeek, Department of Biological Sciences,
Dartmouth College
VI.16 EVOLUTION OF COMMUNITIES
Christine W. Miller, Department of Entomology and
Nematology, University of Florida
VII.5 SEXUAL SELECTION: MALE-MALE COMPETITION
Antónia Monteiro, Department of Ecology and Evolutionary
Biology, Yale University
V.11 EVOLUTION AND DEVELOPMENT: MOLECULES
Jacob A. Moorad, Department of Biology, Duke University
VII.16 AGING AND MENOPAUSE
Allen J. Moore, Department of Genetics, University of
Georgia
VII EVOLUTION OF BEHAVIOR, SOCIETY, AND HUMANS
Patrik Nosil, Department of Ecology and Evolutionary
Biology, University of Colorado, Boulder
VI.9 SPECIATION AND GENOME EVOLUTION
Samir Okasha, Department of Philosophy, University of
Bristol
III.2 UNITS AND LEVELS OF SELECTION
Lorraine Olendzenski, Department of Biology, St. Lawrence
University
II.11 EVOLUTION IN THE PROKARYOTIC GRADE
Kevin E. Omland, Department of Biological Sciences,
University of Maryland, Baltimore County
II.1 INTERPRETATION OF PHYLOGENETIC TREES
H. Allen Orr, Department of Biology, University of
Rochester
VI.8 GENETICS OF SPECIATION
Sarah P. Otto, Department of Zoology, University of British
Columbia
III.9 EVOLUTION OF MODIFIER GENES AND BIOLOGICAL
SYSTEMS
Mark Pagel, School of Biological Sciences, University of
Reading
VIII.9 LINGUISTICS AND THE EVOLUTION OF HUMAN
LANGUAGE
Laura Wegener Parfrey, Department of Chemistry
and Biochemistry, University of Colorado,
Boulder
II.12 ORIGIN AND DIVERSIFICATION OF EUKARYOTES
Bret A. Payseur, Laboratory of Genetics, University of
Wisconsin, Madison
V.13 DISSECTION OF COMPLEX TRAIT EVOLUTION
Contributors xi
Talima Pearson, Department of Biological Sciences,
Northern Arizona University
VIII.4 EVOLUTION AND MICROBIAL FORENSICS
Catherine L. Peichel, Fred Hutchinson Cancer Research
Center, Seattle
V GENES, GENOMES, PHENOTYPES; V.12 GENETICS OF
PHENOTYPIC EVOLUTION
Robert T. Pennock, Lyman Briggs College and Departments
of Philosophy and Computer Science  Engineering,
Michigan State University
VIII.8 EVOLUTION AND COMPUTING
Dmitri A. Petrov, Department of Biology, Stanford
University
V.14 SEARCHING FOR ADAPTATION IN THE GENOME
David W. Pfennig, Department of Biology, University of
North Carolina, Chapel Hill
III.7 RESPONSES TO SELECTION: NATURAL
POPULATIONS
Albert Phillimore, Institute of Evolutionary Biology,
University of Edinburgh
VI.3 GEOGRAPHY, RANGE EVOLUTION, AND SPECIATION
Daniel E. L. Promislow, Department of Pathology,
University of Washington
VII.16 AGING AND MENOPAUSE
Erik Quandt, Department of Chemistry and Biochemistry,
University of Texas at Austin
VIII.7 DIRECTED EVOLUTION
David C. Queller, Department of Biology, Washington
University in St. Louis
III.4 KIN SELECTION AND INCLUSIVE FITNESS; VII.9
COOPERATION AND CONFLICT: MICROBES TO
HUMANS
Bruce Rannala, Department of Evolution and Ecology,
University of California, Davis
II.3 MOLECULAR CLOCK DATING
Richard Ree, Botany Department, Field Museum of Natural
History
II.7 USING PHYLOGENIES TO STUDY PHENOTYPIC
EVOLUTION: COMPARATIVE METHODS AND TESTS OF
ADAPTATION
David Reznick, Department of Biology, University of
California, Riverside
III.11 EVOLUTION OF LIFE HISTORIES
Robert C. Richardson, Department of Philosophy, University
of Cincinnati
VII.12 EVOLUTIONARY PSYCHOLOGY
Ophélie Ronce, Institut des Sciences de l’Evolution,
Université Montpellier 2, Centre National de la Recherche
Scientifique
IV.3 GEOGRAPHIC VARIATION, POPULATION STRUCTURE,
AND MIGRATION
Nick J. Royle, Department of Biosciences, University of
Exeter
VII.8 EVOLUTION OF PARENTAL CARE
Dolph Schluter, Department of Zoology, University of
British Columbia
VI SPECIATION AND MACROEVOLUTION
Eugenie C. Scott, National Center for Science Education, Inc.
VIII.14 CREATIONISM AND INTELLIGENT DESIGN
H. Bradley Shaffer, Department of Ecology and Evolutionary
Biology, University of California, Los Angeles
VIII.6 EVOLUTION AND CONSERVATION
Beth Shapiro, Department of Ecology and Evolutionary
Biology, University of California, Santa Cruz
V.15 ANCIENT DNA
Mark L. Siegal, Department of Biology, New York
University
V.9 EVOLUTION OF MOLECULAR NETWORKS
Per T. Smiseth, Institute of Evolutionary Biology, University
of Edinburgh
VII.8 EVOLUTION OF PARENTAL CARE
Rhonda R. Snook, Department of Animal and Plant Sciences,
University of Sheffield
VII.4 SEXUAL SELECTION AND ITS IMPACT ON MATING
SYSTEMS
Jason E. Stajich, Department of Plant Pathology and
Microbiology, University of California, Riverside
V.3 COMPARATIVE GENOMICS
Stephen C. Stearns, Department of Ecology and Evolutionary
Biology, Yale University
III.1 NATURAL SELECTION, ADAPTATION, AND FITNESS:
OVERVIEW; III.10 EVOLUTION OF REACTION NORMS
Joan E. Strassmann, Department of Biology, Washington
University in St. Louis
III.4 KIN SELECTION AND INCLUSIVE FITNESS; VII.9
COOPERATION AND CONFLICT: MICROBES TO
HUMANS
Sharon Y. Strauss, Department of Ecology and Evolution,
University of California, Davis
III.15 ADAPTATION TO THE BIOTIC ENVIRONMENT
Alan R. Templeton, Department of Biology, Washington
University in St. Louis, Department of Evolutionary and
Environmental Biology, University of Haifa
VIII.11 EVOLUTION AND NOTIONS OF HUMAN RACE;
VIII.12 THE FUTURE OF HUMAN EVOLUTION
John N. Thompson, Department of Ecology and
Evolutionary Biology, University of California, Santa Cruz
VI.7 COEVOLUTION AND SPECIATION
Michelle D. Trautwein, Biodiversity Laboratory, North
Carolina Museum of Natural Sciences
II.16 MAJOR EVENTS IN THE EVOLUTION OF ARTHROPODS
Paul E. Turner, Department of Ecology and Evolutionary
Biology, Yale University
VIII.1 EVOLUTIONARY MEDICINE
Peter C. Wainwright, Department of Evolution and Ecology,
University of California, Davis
III.12 EVOLUTION OF FORM AND FUNCTION
xii Contributors
Michael C. Whitlock, Department of Zoology, University of
British Columbia
I.4 FROM DNA TO PHENOTYPES; IV EVOLUTIONARY
PROCESSES
Brian M. Wiegmann, Department of Entomology, North
Carolina State University
II.16 MAJOR EVENTS IN THE EVOLUTION OF ARTHROPODS
Patricia J. Wittkopp, Department of Ecology and
Evolutionary Biology, University of Michigan
V.7 EVOLUTION OF GENE EXPRESSION
Ziheng Yang, Department of Genetics, Evolution, and
Environment, University College, London
II.3 MOLECULAR CLOCK DATING
Jianzhi Zhang, Department of Ecology and Evolutionary
Biology, University of Michigan
V.5 GENE DUPLICATION
Carl Zimmer, Environmental Studies Program, Yale
University
VIII.15 EVOLUTION AND THE MEDIA
Contributors xiii
I
Introduction
I.1
What Is Evolution?
Jonathan Losos
OUTLINE
1. What is evolution?
2. Evolution: Pattern versus process
3. Evolution: More than changes in the
gene pool
4. In the light of evolution
5. Critiques and the evidence for evolution
6. The pace of evolution
7. Evolution, humans, and society
Evolution refers to change through time as species be-
come modified and diverge to produce multiple descen-
dant species. Evolution and natural selection are often
conflated, but evolution is the historical occurrence of
change, and natural selection is one mechanism—in most
cases the most important—that can cause it. Recent
years have seen a flowering in the field of evolutionary
biology, and much has been learned about the causes and
consequences of evolution. The two main pillars of our
knowledge of evolution come from knowledge of the
historicalrecordofevolutionarychange,deduceddirectly
from the fossil record and inferred from examination of
phylogeny, and from study of the process of evolutionary
change, particularly the effect of natural selection. It is
now apparent that whenselection is strong,evolution can
proceed considerably more rapidly than was generally
envisioned by Darwin. As a result, scientists are realizing
that it is possible to conduct evolutionary experiments in
real time. Recent developments in many areas, including
molecular and developmental biology, have greatly ex-
panded our knowledge and reaffirmed evolution’s central
place in the understanding of biological diversity.
GLOSSARY
Evolution. Descent with modification; transformation
of species through time, including both changes that
occur within species, as well as the origin of new
species.
Natural Selection. The process in which individuals with
a particular trait tend to leave more offspring in the
next generation than do individuals with a different
trait.
Approximately 375 million years ago, a large and vague-
ly salamander-like creature plodded from its aquatic
home and began the vertebrate invasion of land, setting
forththe chain of evolutionary eventsthatled tothe birds
thatfillourskies,thebeaststhatwalkoursoil,mewriting
this chapter, and you reading it. This was, of course, just
one episode in life’s saga: millions of years earlier, plants
had come ashore, followed soon thereafter—or perhaps
simultaneously—byarthropods.Wecouldgobackmuch
earlier, 4 billion years or so, to that fateful day when the
first molecule replicated itself, an important milestone in
the origin of life and the beginning of the evolutionary
pageant. Moving forward, the last few hundred million
years have also had their highs and lows: the origins of
frogs and trees, the end-Permian extinction when 90
percent of all species perished, and the rise and fall of the
dinosaurs.
These vignettes are a few of many waypoints in the
evolutionary chronicle of life on earth. Evolutionary
biologists try to understand this history, explaining how
and why life has taken its particular path. But the study
of evolution involves more than looking backward to try
to understand the past. Evolution is an ongoing process,
one possibly operating at a faster rate now than in times
past in this human-dominated world. Consequently,
evolutionary biology is also forward looking: it includes
the study of evolutionary processes in action today—
how they operate, what they produce—as well as in-
vestigation of how evolution is likely to proceed in the
future. Moreover, evolutionary biology is not solely an
academic matter; evolution affects humans in many
ways, from coping with the emergence of agricultural
pests and disease-causing organisms to understanding
the workings of our own genome. Indeed, evolutionary
science hasbroad relevance,playingan importantrole in
advances in many areas, from computer programming
to medicine to engineering.
1. WHAT IS EVOLUTION?
Lookuptheword“evolution”intheonlineversionofthe
Oxford English Dictionary, and you will find 11 defini-
tions and numerous subdefinitions, ranging from math-
ematical (“the successive transformation of a curve by
the alteration of the conditionswhich defineit”) tochem-
ical (“the emission or release of gas, heat, light, etc.”) to
military (“a manoeuvre executed by troops or ships to
adopt a different tactical formation”). Even with ref-
erence to biology, there are several definitions, including
“emergence or release from an envelope or enclosing
structure; (also) protrusion, evagination,” not to men-
tion “rare” and “historical” usage related to the concept
of preformation of embryos. Even among evolutionary
biologists, evolution is defined in different ways. For
example, one widely read textbook refers to evolution as
“changes in the properties of groups of organisms over
the course of generations” (Futuyma 2005), whereas
another defines it as “changes in allele frequencies over
time” (Freeman and Herron 2007).
One might think that—as in so many other areas of
evolutionary biology—we could look to Darwin for
clarity. But in the first edition of On the Origin of Spe-
cies, the term “evolution” never appears (though the last
word of the book is “evolved”); not until the sixth edi-
tion does Darwin use “evolution.” Rather, Darwin’s
term of choice is “descent with modification,” a simple
phrase that captures the essence of what evolutionary
biology is all about: the study of the transformation of
species through time, including both changes that occur
within species, as well as the origin of new species.
2. EVOLUTION: PATTERN VERSUS PROCESS
Many people—sometimes even biologists—equate evo-
lution with natural selection, but the two are not the
same. Natural selection is one process that can cause
evolutionary change, but natural selection can occur
without producing evolutionary change. Conversely,
processes other than natural selection can lead to
evolution.
Natural selection within populations refers to the sit-
uation in which individuals with one variant of a trait
(say, blue eyes) tend to leave more offspring that are
healthy and fertile in the next generation than do in-
dividuals with an alternative variant of the trait. Such
selection can occur in many ways, for example, if the
variantleadstogreaterlongevity,greaterattractivenessto
members of the other sex, or greater number of offspring
per breeding event. The logic behind natural selection is
unassailable. If some trait variant is causally related to
greater reproductive success, then more members of the
population will have that variant in the next generation;
continued over many generations, such selection can
greatly change the constitution of a population.
But there is a catch. Natural selection can occur with-
out leading to evolution if differences among individuals
are not genetically based. For natural selection to cause
evolutionary change, trait variants must be transmitted
from parent to offspring; if that is the case, then offspring
will resemble their parents and the trait variants possessed
by the parents that produce the most offspring will in-
crease in frequency in the next generation.
However, offspring do not always resemble their
parents. In some cases, individuals vary phenotypically
not because they are different genetically, but because
they experienced different environments during growth
(this is the “nurture” part of the nature versus nurture
debate; see chapters III.10 and VII.1). If, in fact, varia-
tion in a population is not genetically based, then se-
lection will have no evolutionary consequence; in-
dividuals surviving and producing many offspring will
not differ genetically from those that fail to prosper, and
as a result, the gene pool of the population will not
change. Nonetheless, much of the phenotypic variation
within a population is, in fact, genetically based; con-
sequently, natural selection often does lead to evolu-
tionary change.
But that does not mean that the occurrence of evo-
lutionary change necessarily implies the action of nat-
ural selection. Other processes—especially mutation,
genetic drift, and immigration of individuals with dif-
ferent genetic constitutions—also can cause a change in
the genetic makeup of a population from one generation
to the next (see Section IV: Evolutionary Processes). In
other words, natural selection can cause adaptive evo-
lutionary change, but not all evolution is adaptive.
These caveats notwithstanding, 150 years of research
have made clear that naturalselectionisa powerful force
responsible for much of the significant evolutionary
change that has occurred over the history of life. As the
chapters in Section II: Phylogenetics and the History of
Life, and Section III: Natural Selection and Adaptation,
demonstrate, natural selection can operate in many
ways, and scientists have correspondingly devised many
methods to detect it, both through studies of the phe-
notype and of DNA itself (see also chapter V.14).
3. EVOLUTION: MORE THAN CHANGES IN THE
GENE POOL
During the heyday of population genetics in the middle
decades of the last century, many biologists equated
4 Introduction
evolution with changes from one generation to the next
in gene frequencies (gene frequency refers to the fre-
quencies of different alleles of a gene; for background
on genetic variation, see chapter I.4). The “Modern
Synthesis” of the 1930s and 1940s led to several decades
in which the field was primarily concerned with the ge-
netics of populations with an emphasis on natural se-
lection (see chapter I.2). This focus was sharpened by the
advent of molecular approaches to studying evolution.
Starting in 1960 with the application of enzyme elec-
trophoresis techniques, biologists could, for the first
time, directly assess the extentof genetic variation within
populations. To everyone’s surprise, populations were
found to contain much more variation than expected.
This finding both challenged the view that natural se-
lection was the dominant force guiding evolutionary
change (see discussion of “neutralists” in chapters I.2
and V.1), yet further directed attention to the genetics of
populations. With more advanced molecular techniques
available today, the situation has not changed. There is
much more variation than we first suspected.
The last 35 years have seen a broadening of evolu-
tionary inquiry as the field has recognized that there is
more to understanding evolutionary change than study-
ing what happens to genes within populations—though
this area remains a critically important part of evolu-
tionary inquiry. Three aspects of expansion in evolu-
tionary thinking are particularly important.
First, phenotypic evolution results from evolutionary
change in the developmental process that transforms a
single-celled fertilized egg into an adult organism. Al-
though under genetic control, development is an in-
tricate process that cannot be understood by examina-
tion of DNA sequences alone. Rather, understanding
how phenotypes evolve, and the extent to which devel-
opmental systems constrain and direct evolutionary
change, requires detailed molecular and embryological
knowledge (see chapters V.10 and V.11).
Second, history is integral to understanding evolution
(see introduction to Section II: Phylogenetics and the
History of Life). The study of fossils—paleontology—
provides the primary, almost exclusive, direct evidence
of life in the past. Somewhat moribund in the middle of
the last century, paleontology has experienced a resur-
gence in recent decades owing to both dramatic new dis-
coveries stemming from an upsurge in paleontological
exploration, and new ideas about evolution inspired by
and primarily testable with fossil data, such as theories
concerning punctuated equilibrium and stasis, species
selection, and mass extinction. Initially critical in the
development and acceptance of evolutionary theory,
paleontology has once again become an important and
vibrant part of evolutionary biology (see chapter II.9 and
others in Section II).
Concurrently, a more fundamental revolution em-
phasizing the historical perspective has taken place over
the last 30 years with the realization that information on
phylogenetic relationships—that is, the tree of life, the
pattern of descent and relationship among species—is
critical in interpreting all aspects of evolution above the
population level. Beginning with a transformation in the
field of systematics concerning how phylogenetic re-
lationships are inferred, this “tree-thinking” approach
now guides study not only of all aspects of macroevolu-
tion but also of many population-level phenomena.
Finally, life is hierarchically organized. Genes are lo-
cated within individuals, individuals within populations,
populations within species, and species within clades
(a clade consists of an ancestral species and all its de-
scendants). Population genetics concerns what happens
among individuals within a population, but evolutionary
change can occur at all levels. For example, why are there
more than 2000 species of rodents but only 3 species of
monotremes (the platypus and echidnas), a much older
clade of mammals? One cannot look at questions con-
cerning natural selection within a population to answer
this question. Rather, one must inquire about properties
of entire species. Is there some attribute of rodents that
makes them particularly prone to speciate or to avoid
extinction? Similarly, why is there so much seemingly
useless noncoding DNA in the genomes of many species
(see chapter V.2)? One possibility is that some genes are
particularly adept at mutating to multiply the number of
copies of that gene within a genome; such DNA might
increase in frequency in the genome even if such multi-
plication has no benefit to the individual in whose body
the DNA resides. Just as selection among individual or-
ganisms on heritable traits can lead to evolutionary
change within populations, selection among entities at
other levels (species, genes) can also lead to evolutionary
change, as long as those entities have traits that are
transmitted to their offspring (be they descendant species
or genes) and affect the number of descendants they pro-
duce.Theupshotisthatevolutionoccursatmultiplelevels
of the hierarchy of life; to understand its rich complexity
we must study evolution at these distinct levels as well as
theinteractionsamongthem.Whathappens,forexample,
when a trait that benefits an individual within a popula-
tion (perhaps cannibalism—more food, fewer competi-
tors!) has detrimental effects at the level of species?
Although evolutionary biology has expanded in
scope, genetic change is still its fundamental foundation.
Nonetheless, in recent years attention has focused on
variation that is not genetically based. Phenotypic plas-
ticity—the ability of a single genotype to produce dif-
ferent phenotypes when exposed to different environ-
ments—may itself be adaptive (see chapter III.10). If
individuals in a population are likely to experience
What Is Evolution? 5
differentconditionsastheydevelop,thentheevolutionof
a genotype that could produce appropriate phenotypes
depending on circumstances would be advantageous.
Although selection on these different phenotypes would
not lead to evolutionary change, the degree of plasticity
itself can evolve if differences in extent of plasticity lead
to differences in the number of surviving offspring. In-
deed, an open question is, why don’t populations evolve
to become infinitely malleable, capable of producing the
appropriatephenotypeforanyenvironment?Presumably,
plasticity has an associated cost such that adaptation to
different environments often occurs by genetic differ-
entiation rather than by the evolution of a single genotype
that can produce different phenotypes. Such costs, how-
ever, have proven difficult to demonstrate.
Differences observed among populations may also
reflect plastic responses to different environmental con-
ditions and thus may not reflect genetic differentiation.
However, if consistently transmitted from one genera-
tion to the next, such nongenetic differences may lead to
divergent selective pressures on traits that are genetically
determined, thus promoting evolutionary divergence
between the populations. One particular example con-
cerns behavior, which is highly variable in response to
the environment—an extreme manifestation of plastic-
ity (see chapter VIII.10). Learned behaviors that are
transmitted from one generation to the next—often
called traditions or culture—occur not only in humans
but in other animals, not only our near relatives the apes
but also cetaceans, birds, and others. Such behavioral
differences among populations would not reflect genetic
differentiation, but they might set the stage for genetic
divergence in traits relating to the behaviors. One can
easily envision, for example, how chimpanzee popula-
tions that use different tools—such as delicate twigs to
probe termite mounds, or heavy stones to pound nuts—
might evolve different morphological features to en-
hance the effectiveness of these behaviors. A concrete
example involves human populations that tend cattle—
surely a nongenetically based behavior—and have
evolved genetic changes to permit the digestion of milk
in adults.
4. IN THE LIGHT OF EVOLUTION
In a 1964 address to the American Society of Zoologists,
the distinguished Russian-born biologist Theodosius
Dobzhansky proclaimed “nothing makes sense in biol-
ogy except in the light of evolution.” Ever since, evolu-
tionary biologists have trotted out this phrase (or some
permutation of it) to emphasize the centrality of evolu-
tion in understanding the biological world. Nonetheless,
for much of the twentieth century, the pervasive im-
portance of an evolutionary perspective was not at all
obvious to many biologists, some of whom considered
Dobzhansky’s claim to be self-serving hype. One could
argue, for example, that the enormous growth in our
understanding of molecular biology from 1950 to 2000
was made with little involvement or insight from evo-
lutionary biology. Indeed, to the practicing molecular
biologist in the 1980s and 1990s, evolutionary biology
was mostly irrelevant.
Now, nothing could be further from the truth. When
results of the human genome sequencing project first
appeared in2000, many initiallybelieved that a thorough
understanding of human biology would soon follow,
answering questions about the genetic basis of human
diseases and phenotypic variation among individuals.
These hopes were quickly dashed—the genetic code,after
all, is nothing more than a long list of letters (A, C, G, and
T, the abbreviations of the four nucleotide building
blocks of DNA). Much of the genome of many species
seems to have no function and is just, in some sense,
functionless filler; as a result, picking out where the genes
lieinthis4billion–longstringofalphabetspaghetti,much
less figuring out how these genes function, is not easy.
Sowhere did molecular biologists turn? Tothe fieldof
evolutionary biology! Genomicists soon realized that the
best way to understand the human genome was to study
it in the context of its evolutionary history, by comparing
human sequences with those of other species in a phy-
logenetic framework. One method for locating genes,
for example, is to examine comparable parts of the
genome of different species. The underlying rationale is
that genes evolve more slowly than other parts of the
genome. Specifically, nonfunctioning stretches of DNA
tend to evolve differences through time as random mu-
tations become established (the process of genetic drift;
see chapter IV.1), but functioning genes tend to diverge
less, because natural selection removes deleterious mu-
tations when they arise, keeping the DNA sequence sim-
ilar among species. As a result, examination of the
amount of divergence between two species relative to the
amountof time since theyshareda commonancestor can
pinpoint stretches of DNA where evolution has occurred
slowly, thus identifying the position of functional genes.
Moreover, how a gene functions can often be deduced by
comparing its function with that of homologous genes in
other species and using a phylogeny to reconstruct the
gene’s evolutionary history (see chapter V.14).
And thus was born the effort to sequence the ge-
nomes of other species (see chapter V.3). At first, the
nascent field of comparative genomics focused on pri-
mates and model laboratory species such as mice and
fruit flies, the former to permit comparisons of the
human genome with that of our close evolutionary rel-
atives, the latter to take advantage of the great under-
standing of the genomic systems of well-studied species.
6 Introduction
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CHAPTER XXIV.
THE REASON AND END OF DIVINE PUNISHMENTS.
Now that is in our power, of which equally with its opposite we
are masters,—as, say, to philosophize or not, to believe or
disbelieve. In consequence, then, of our being equally masters of
each of the opposites, what depends on us is found possible. Now
the commandments may be done or not done by us, who, as is
reasonable, are liable to praise and blame. And those, again, who
are punished on account of sins committed by them, are punished
for them alone; for what is done is past, and what is done can never
be undone. The sins committed before faith are accordingly forgiven
by the Lord, not that they may be undone, but as if they had not
been done. “But not all,” says Basilides, “but only sins involuntary
and in ignorance, are forgiven;” as would be the case were it a man,
and not God, that conferred such a boon. To such an one Scripture
says, “Thou thoughtest that I would be like thee.”[662] But if we are
punished for voluntary sins, we are punished not that the sins which
are done may be undone, but because they were done. But
punishment does not avail to him who has sinned, to undo his sin,
but that he may sin no more, and that no one else fall into the like.
Therefore the good God corrects for these three causes: First, that
he who is corrected may become better than his former self; then
that those who are capable of being saved by examples may be
driven back, being admonished; and thirdly, that he who is injured
may not be readily despised, and be apt to receive injury. And there
are two methods of correction—the instructive and the punitive,
which we have called the disciplinary. It ought to be known, then,
that those who fall into sin after baptism[663] are those who are
subjected to discipline; for the deeds done before are remitted, and
those done after are purged. It is in reference to the unbelieving
that it is said, “that they are reckoned as the chaff which the wind
drives from the face of the earth, and the drop which falls from a
vessel.”[664]
CHAPTER XXV.
TRUE PERFECTION CONSISTS IN THE KNOWLEDGE AND LOVE OF GOD.
“Happy he who possesses the culture of knowledge, and is not
moved to the injury of the citizens or to wrong actions, but
contemplates the undecaying order of immortal nature, how and in
what way and manner it subsists. To such the practice of base deeds
attaches not.” Rightly, then, Plato says, “that the man who devotes
himself to the contemplation of ideas will live as a god among men;
now the mind is the place of ideas, and God is mind.” He says that
he who contemplates the unseen God lives as a god among men.
And in the Sophist, Socrates calls the stranger of Elea, who was a
dialectician, “god:” “Such are the gods who, like stranger guests,
frequent cities. For when the soul, rising above the sphere of
generation, is by itself apart, and dwells amidst ideas,” like the
Coryphæus in Theætetus, now become as an angel, it will be with
Christ, being rapt in contemplation, ever keeping in view the will of
God; in reality
“Alone wise, while these flit like shadows.”[665]
“For the dead bury their dead.” Whence Jeremiah says: “I will fill it
with the earth-born dead whom mine anger has smitten.”[666]
God, then, being not a subject for demonstration, cannot be the
object of science. But the Son is wisdom, and knowledge, and truth,
and all else that has affinity thereto. He is also susceptible of
demonstration and of description. And all the powers of the Spirit,
becoming collectively one thing, terminate in the same point—that
is, in the Son. But He is incapable of being declared, in respect of
the idea of each one of His powers. And the Son is neither simply
one thing as one thing, nor many things as parts, but one thing as
all things; whence also He is all things. For He is the circle of all
powers rolled and united into one unity. Wherefore the Word is
called the Alpha and the Omega, of whom alone the end becomes
beginning, and ends again at the original beginning without any
break. Wherefore also to believe in Him, and by Him, is to become a
unit, being indissolubly united in Him; and to disbelieve is to be
separated, disjoined, divided.
“Wherefore thus saith the Lord, Every alien son is uncircumcised
in heart, and uncircumcised in flesh” (that is, unclean in body and
soul): “there shall not enter one of the strangers into the midst of
the house of Israel, but the Levites.”[667] He calls those that would
not believe, but would disbelieve, strangers. Only those who live
purely being true priests of God. Wherefore, of all the circumcised
tribes, those anointed to be high priests, and kings, and prophets,
were reckoned more holy. Whence He commands them not to touch
dead bodies, or approach the dead; not that the body was polluted,
but that sin and disobedience were incarnate, and embodied, and
dead, and therefore abominable. It was only, then, when a father
and mother, a son and daughter died, that the priest was allowed to
enter, because these were related only by flesh and seed, to whom
the priest was indebted for the immediate cause of his entrance into
life. And they purify themselves seven days, the period in which
Creation was consummated. For on the seventh day the rest is
celebrated; and on the eighth he brings a propitiation, as is written
in Ezekiel, according to which propitiation the promise is to be
received.[668] And the perfect propitiation, I take it, is that propitious
faith in the gospel which is by the law and the prophets, and the
purity which shows itself in universal obedience, with the
abandonment of the things of the world; in order to that grateful
surrender of the tabernacle, which results from the enjoyment of the
soul. Whether, then, the time be that which through the seven
periods enumerated returns to the chiefest rest,[669] or the seven
heavens, which some reckon one above the other; or whether also
the fixed sphere which borders on the intellectual world be called the
eighth, the expression denotes that the Gnostic ought to rise out of
the sphere of creation and of sin. After these seven days, sacrifices
are offered for sins. For there is still fear of change, and it touches
the seventh circle. The righteous Job says: “Naked came I out of my
mother’s womb, and naked shall I return there;”[670] not naked of
possessions, for that were a trivial and common thing; but, as a just
man, he departs naked of evil and sin, and of the unsightly shape
which follows those who have led bad lives. For this was what was
said, “Unless ye be converted, and become as children,”[671] pure in
flesh, holy in soul by abstinence from evil deeds; showing that He
would have us to be such as also He generated us from our mother
—the water.[672] For the intent of one generation succeeding
another is to immortalize by progress. “But the lamp of the wicked
shall be put out.”[673] That purity in body and soul which the Gnostic
partakes of, the all-wise Moses indicated, by employing repetition in
describing the incorruptibility of body and of soul in the person of
Rebecca, thus: “Now the virgin was fair, and man had not known
her.”[674] And Rebecca, interpreted, means “glory of God;” and the
glory of God is immortality. This is in reality righteousness, not to
desire other things, but to be entirely the consecrated temple of the
Lord. Righteousness is peace of life and a well-conditioned state, to
which the Lord dismissed her when He said, “Depart into
peace.”[675] For Salem is, by interpretation, peace; of which our
Saviour is enrolled King, as Moses says, Melchizedek king of Salem,
priest of the most high God, who gave bread and wine, furnishing
consecrated food for a type of the Eucharist. And Melchizedek is
interpreted “righteous king;” and the name is a synonym for
righteousness and peace. Basilides, however, supposes that
Righteousness and her daughter Peace dwell stationed in the eighth
sphere.
But we must pass from physics to ethics, which are clearer; for
the discourse concerning these will follow after the treatise in hand.
The Saviour Himself, then, plainly initiates us into the mysteries,
according to the words of the tragedy:[676]
“Seeing those who see, he also gives the orgies.”
And if you ask,
“These orgies, what is their nature?”
You will hear again:
“It is forbidden to mortals uninitiated in the Bacchic rites to know.”
And if any one will inquire curiously what they are, let him hear:
“It is not lawful for thee to hear, but they are worth knowing;
The rites of the God detest him who practises impiety.”
Now God, who is without beginning, is the perfect beginning of the
universe, and the producer of the beginning. As, then, He is being,
He is the first principle of the department of action, as He is good, of
morals; as He is mind, on the other hand, He is the first principle of
reasoning and of judgment. Whence also He alone is Teacher, who is
the only Son of the Most High Father, the Instructor of men.
CHAPTER XXVI.
HOW THE PERFECT MAN TREATS THE BODY AND THE THINGS OF THE WORLD.
Those, then, who run down created existence and vilify the body
are wrong; not considering that the frame of man was formed erect
for the contemplation of heaven, and that the organization of the
senses tends to knowledge; and that the members and parts are
arranged for good, not for pleasure. Whence this abode becomes
receptive of the soul which is most precious to God; and is dignified
with the Holy Spirit through the sanctification of soul and body,
perfected with the perfection of the Saviour. And the succession of
the three virtues is found in the Gnostic, who morally, physically, and
logically occupies himself with God. For wisdom is the knowledge of
things divine and human; and righteousness is the concord of the
parts of the soul; and holiness is the service of God. But if one were
to say that he disparaged the flesh, and generation on account of it,
by quoting Isaiah, who says, “All flesh is grass, and all the glory of
man as the flower of grass: the grass is withered, and the flower has
fallen; but the word of the Lord endureth for ever;”[677] let him hear
the Spirit interpreting the matter in question by Jeremiah, “And I
scattered them like dry sticks, that are made to fly by the wind into
the desert. This is the lot and portion of your disobedience, saith the
Lord. As thou hast forgotten me, and hast trusted in lies, so will I
discover thy hinder parts to thy face; and thy disgrace shall be seen,
thy adultery, and thy neighing,” and so on.[678] For “the flower of
grass,” and “walking after the flesh,” and “being carnal,” according to
the apostle, are those who are in their sins. The soul of man is
confessedly the better part of man, and the body the inferior. But
neither is the soul good by nature, nor, on the other hand, is the
body bad by nature. Nor is that which is not good straightway bad.
For there are things which occupy a middle place, and among them
are things to be preferred, and things to be rejected. The
constitution of man, then, which has its place among things of
sense, was necessarily composed of things diverse, but not opposite
—body and soul.
Always therefore the good actions, as better, attach to the better
and ruling spirit; and voluptuous and sinful actions are attributed to
the worse, the sinful one.
Now the soul of the wise man and Gnostic, as sojourning in the
body, conducts itself towards it gravely and respectfully, not with
inordinate affections, as about to leave the tabernacle if the time of
departure summon. “I am a stranger in the earth, and a sojourner
with you,” it is said.[679] And hence Basilides says, that he
apprehends that the election are strangers to the world, being
supramundane by nature. But this is not the case. For all things are
of one God. And no one is a stranger to the world by nature, their
essence being one, and God one. But the elect man dwells as a
sojourner, knowing all things to be possessed and disposed of; and
he makes use of the things which the Pythagoreans make out to be
the threefold good things. The body, too, as one sent on a distant
pilgrimage, uses inns and dwellings by the way, having care of the
things of the world, of the places where he halts; but leaving his
dwelling-place and property without excessive emotion; readily
following him that leads him away from life; by no means and on no
occasion turning back; giving thanks for his sojourn, and blessing
[God] for his departure, embracing the mansion that is in heaven.
“For we know, that, if the earthly house of our tabernacle be
dissolved, we have a building of God, an house not made with
hands, eternal in the heavens. For we that are in this tabernacle do
groan, desiring to be clothed upon with our house which is from
heaven: if so be that being clothed we shall not be found naked. For
we walk by faith, not by sight,”[680] as the apostle says; “and we are
willing rather to be absent from the body, and present with God.”
The rather is in comparison. And comparison obtains in the case of
things that fall under resemblance; as the more valiant man is more
valiant among the valiant, and most valiant among cowards. Whence
he adds, “Wherefore we strive, whether present or absent, to be
accepted with Him,”[681] that is, God, whose work and creation are
all things, both the world and things supramundane. I admire
Epicharmus, who clearly says:
“Endowed with pious mind, you will not, in dying,
Suffer aught evil. The spirit will dwell in heaven above;”
and the minstrel[682] who sings:
“The souls of the wicked flit about below the skies on earth,
In murderous pains beneath inevitable yokes of evils;
But those of the pious dwell in the heavens,
Hymning in songs the Great, the Blessed One.”
The soul is not then sent down from heaven to what is worse. For
God works all things up to what is better. But the soul which has
chosen the best life—the life that is from God and righteousness—
exchanges earth for heaven. With reason therefore, Job, who had
attained to knowledge, said, “Now I know that Thou canst do all
things; and nothing is impossible to Thee. For who tells me of what I
know not, great and wonderful things with which I was
unacquainted? And I felt myself vile, considering myself to be earth
and ashes.”[683] For he who, being in a state of ignorance, is sinful,
“is earth and ashes;” while he who is in a state of knowledge, being
assimilated as far as possible to God, is already spiritual, and so
elect. And that Scripture calls the senseless and disobedient “earth,”
will be made clear by Jeremiah the prophet, saying, in reference to
Joachim and his brethren, “Earth, earth, hear the word of the Lord;
Write this man, as man excommunicated.”[684] And another prophet
says again, “Hear, O heaven; and give ear, O earth,”[685] calling
understanding “ear,” and the soul of the Gnostic, that of the man
who has applied himself to the contemplation of heaven and divine
things, and in this way has become an Israelite, “heaven.” For again
he calls him who has made ignorance and hardness of heart his
choice, “earth.” And the expression “give ear” he derives from the
“organs of hearing,” “the ears,” attributing carnal things to those
who cleave to the things of sense. Such are they of whom Micah the
prophet says, “Hear the word of the Lord, ye peoples who dwell with
pangs.”[686] And Abraham said, “By no means. The Lord is He who
judgeth the earth;”[687] “since he that believeth not, is,” according to
the utterance of the Saviour, “condemned already.”[688] And there is
written in the Kings[689] the judgment and sentence of the Lord,
which stands thus: “The Lord hears the righteous, but the wicked He
saveth not, because they do not desire to know God.” For the
Almighty will not accomplish what is absurd. What do the heresies
say to this utterance, seeing Scripture proclaims the Almighty God to
be good, and not the author of evil and wrong, if indeed ignorance
arises from one not knowing? But God does nothing absurd. “For
this God,” it is said, “is our God, and there is none to save besides
Him.”[690] “For there is no unrighteousness with God,”[691] according
to the apostle. And clearly yet the prophet teaches the will of God,
and the gnostic proficiency, in these words: “And now, Israel, what
doth the Lord God require of thee, but to fear the Lord thy God, and
walk in all His ways, and love Him, and serve Him alone?”[692] He
asks of thee, who hast the power of choosing salvation. What is it,
then, that the Pythagoreans mean when they bid us “pray with the
voice?” As seems to me, not that they thought the Divinity could not
hear those who speak silently, but because they wished prayers to
be right, which no one would be ashamed to make in the knowledge
of many. We shall, however, treat of prayer in due course by and by.
But we ought to have works that cry aloud, as becoming “those who
walk in the day.”[693] “Let thy works shine,”[694] and behold a man
and his works before his face. “For behold God and His works.”[695]
For the gnostic must, as far as is possible, imitate God. And the
poets call the elect in their pages godlike and gods, and equal to the
gods, and equal in sagacity to Zeus, and having counsels like the
gods, and resembling the gods,—nibbling, as seems to me, at the
expression, “in the image and likeness.”[696]
Euripides accordingly says, “Golden wings are round my back, and
I am shod with the winged sandals of the Sirens; and I shall go aloft
into the wide ether, to hold converse with Zeus.”
But I shall pray the Spirit of Christ to wing me to my Jerusalem.
For the Stoics say that heaven is properly a city, but places here on
earth are not cities; for they are called so, but are not. For a city is
an important thing, and the people a decorous body, and a multitude
of men regulated by law as the church by the word—a city on earth
impregnable—free from tyranny; a product of the divine will on earth
as in heaven. Images of this city the poets create with their pen. For
the Hyperboreans, and the Arimaspian cities, and the Elysian plains,
are commonwealths of just men. And we know Plato’s city placed as
a pattern in heaven.
BOOK V.
CHAPTER I.
ON FAITH.
f the gnostic so much has been cursorily, as it were,
written. We proceed now to the sequel, and must again
contemplate faith; for there are some that draw the
distinction, that faith has reference to the Son, and
knowledge to the Spirit. But it has escaped their notice
that, in order to believe truly in the Son, we must believe that He is
the Son, and that He came, and how, and for what, and respecting
His passion; and we must know who is the Son of God. Now neither
is knowledge without faith, nor faith without knowledge. Nor is the
Father without the Son; for the Son is with the Father. And the Son
is the true teacher respecting the Father; and that we may believe in
the Son, we must know the Father, with whom also is the Son.
Again, in order that we may know the Father, we must believe in the
Son, that it is the Son of God who teaches; for from faith to
knowledge by the Son is the Father. And the knowledge of the Son
and Father, which is according to the gnostic rule—that which in
reality is gnostic—is the attainment and comprehension of the truth
by the truth.
We, then, are those who are believers in what is not believed, and
who are gnostics as to what is unknown; that is, gnostics as to what
is unknown and disbelieved by all, but believed and known by a few;
and gnostics, not describing actions by speech, but gnostics in the
exercise of contemplation. Happy is he who speaks in the ears of the
hearing. Now Faith is the ear of the soul. And such the Lord
intimates faith to be, when He says, “He that hath ears to hear, let
him hear;”[697] so that by believing he may comprehend what He
says, as He says it. Homer, too, the oldest of the poets, using the
word “hear” instead of “perceive”—the specific for the generic term
—writes:
“Him most they heard.”[698]
For, in fine, the agreement and harmony of the faith of both[699]
contribute to one end—salvation. We have in the apostle an unerring
witness: “For I desire to see you, that I may impart unto you some
spiritual gift, in order that ye may be strengthened; that is, that I
may be comforted in you, by the mutual faith of you and me.”[700]
And further on again he adds, “The righteousness of God is revealed
from faith to faith.”[701] The apostle, then, manifestly announces a
double faith, or rather one which admits of growth and perfection;
for the common faith lies beneath as a foundation. To those,
therefore, who desire to be healed, and are moved by faith, He
added, “Thy faith hath saved thee.”[702] But that which is excellently
built upon is consummated in the believer, and is again perfected by
the faith which results from instruction and the word, in order to the
performance of the commandments. Such were the apostles, in
whose case it is said that “faith removed mountains and transplanted
trees.”[703] Whence, perceiving the greatness of its power, they
asked “that faith might be added to them;”[704] a faith which
salutarily bites the soil “like a grain of mustard,” and grows
magnificently in it, to such a degree that the reasons of things
sublime rest on it. For if one by nature knows God, as Basilides
thinks, who calls intelligence of a superior order at once faith and
kingship, and a creation worthy of the essence of the Creator; and
explains that near Him exists not power, but essence and nature and
substance; and says that faith is not the rational assent of the soul
exercising free-will, but an undefined beauty, belonging immediately
to the creature;—the precepts both of the Old and of the New
Testament are, then, superfluous, if one is saved by nature, as
Valentinus would have it, and is a believer and an elect man by
nature, as Basilides thinks; and nature would have been able, one
time or other, to have shone forth, apart from the Saviour’s
appearance. But were they to say that the visit of the Saviour was
necessary, then the properties of nature are gone from them, the
elect being saved by instruction, and purification, and the doing of
good works. Abraham, accordingly, who through hearing believed
the voice, which promised under the oak in Mamre, “I will give this
land to thee, and to thy seed,” was either elect or not. But if he was
not, how did he straightway believe, as it were naturally? And if he
was elect, their hypothesis is done away with, inasmuch as even
previous to the coming of the Lord an election was found, and that
saved: “For it was reckoned to him for righteousness.”[705] For if any
one, following Marcion, should dare to say that the Creator
(Δημιουργόν) saved the man that believed on him, even before the
advent of the Lord, (the election being saved with their own proper
salvation); the power of the good Being will be eclipsed; inasmuch
as late only, and subsequent to the Creator spoken of by them in
words of good omen, it made the attempt to save, and by his
instruction, and in imitation of him. But if, being such, the good
Being save, according to them; neither is it his own that he saves,
nor is it with the consent of him who formed the creation that he
essays salvation, but by force or fraud. And how can he any more be
good, acting thus, and being posterior? But if the locality is different,
and the dwelling-place of the Omnipotent is remote from the
dwelling-place of the good God; yet the will of him who saves,
having been the first to begin, is not inferior to that of the good
God. From what has been previously proved, those who believe not
are proved senseless: “For their paths are perverted, and they know
not peace,” saith the prophet.[706] “But foolish and unlearned
questions” the divine Paul exhorted to “avoid, because they gender
strifes.”[707] And Æschylus exclaims:
“In what profits not, labour not in vain.”
For that investigation, which accords with faith, which builds, on the
foundation of faith, the august knowledge of the truth, we know to
be the best. Now we know that neither things which are clear are
made subjects of investigation, such as if it is day, while it is day;
nor things unknown, and never destined to become clear, as
whether the stars are even or odd in number; nor things convertible;
and those are so which can be said equally by those who take the
opposite side, as if what is in the womb is a living creature or not. A
fourth mode is, when, from either side of those, there is advanced
an unanswerable and irrefragable argument. If, then, the ground of
inquiry, according to all of these modes, is removed, faith is
established. For we advance to them the unanswerable
consideration, that it is God who speaks and comes to our help in
writing, respecting each one of the points regarding which I
investigate. Who, then, is so impious as to disbelieve God, and to
demand proofs from God as from men? Again, some questions
demand the evidence of the senses, as if one were to ask whether
the fire be warm, or the snow white; and some admonition and
rebuke, as the question if you ought to honour your parents. And
there are those that deserve punishment, as to ask proofs of the
existence of Providence. There being then a Providence, it were
impious to think that the whole of prophecy and the economy in
reference to a Saviour did not take place in accordance with
Providence. And perchance one should not even attempt to
demonstrate such points, the divine Providence being evident from
the sight of all its skilful and wise works which are seen, some of
which take place in order, and some appear in order. And He who
communicated to us being and life, has communicated to us also
reason, wishing us to live rationally and rightly. For the Word of the
Father of the universe is not the uttered word (λόγος προφορικός),
but the wisdom and most manifest kindness of God, and His power
too, which is almighty and truly divine, and not incapable of being
conceived by those who do not confess—the all-potent will. But
since some are unbelieving, and some are disputatious, all do not
attain to the perfection of the good. For neither is it possible to
attain it without the exercise of free choice; nor does the whole
depend on our own purpose; as, for example, what is destined to
happen. “For by grace we are saved:” not, indeed, without good
works; but we must, by being formed for what is good, acquire an
inclination for it. And we must possess the healthy mind which is
fixed on the pursuit of the good; in order to which we have the
greatest need of divine grace, and of right teaching, and of holy
susceptibility, and of the drawing of the Father to Him. For, bound in
this earthly body, we apprehend the objects of sense by means of
the body; but we grasp intellectual objects by means of the logical
faculty itself. But if one expect to apprehend all things by the senses,
he has fallen far from the truth. Spiritually, therefore, the apostle
writes respecting the knowledge of God, “For now we see as
through a glass, but then face to face.”[708] For the vision of the
truth is given but to few. Accordingly, Plato says in the Epinomis, “I
do not say that it is possible for all to be blessed and happy; only a
few. Whilst we live, I pronounce this to be the case. But there is a
good hope that after death I shall attain all.” To the same effect is
what we find in Moses: “No man shall see my face, and live.”[709]
For it is evident that no one during the period of life has been able
to apprehend God clearly. But “the pure in heart shall see God,”[710]
when they arrive at the final perfection. For since the soul became
too enfeebled for the apprehension of realities, we needed a divine
teacher. The Saviour is sent down—a teacher and leader in the
acquisition of the good—the secret and sacred token of the great
Providence. “Where, then, is the scribe? where is the searcher of this
world? Hath not God made foolish the wisdom of this world?”[711] it
is said. And again, “I will destroy the wisdom of the wise, and bring
to nothing the understanding of the prudent,”[712] plainly of those
wise in their own eyes, and disputatious. Excellently therefore
Jeremiah says, “Thus saith the Lord, Stand in the ways, and ask for
the eternal paths, what is the good way, and walk in it, and ye shall
find expiation for your souls.”[713] Ask, he says, and inquire of those
who know, without contention and dispute. And on learning the way
of truth, let us walk on the right way, without turning till we attain to
what we desire. It was therefore with reason that the king of the
Romans (his name was Numa), being a Pythagorean, first of all men,
erected a temple to Faith and Peace. “And to Abraham, on believing,
righteousness was reckoned.”[714] He, prosecuting the lofty
philosophy of aerial phenomena, and the sublime philosophy of the
movements in the heavens, was called Abram, which is interpreted
“sublime father.”[715] But afterwards, on looking up to heaven,
whether it was that he saw the Son in the spirit, as some explain, or
a glorious angel, or in any other way recognised God to be superior
to the creation, and all the order in it, he receives in addition the
Alpha, the knowledge of the one and only God, and is called
Abraam, having, instead of a natural philosopher, become wise, and
a lover of God. For it is interpreted, “elect father of sound.” For by
sound is the uttered word: the mind is its father; and the mind of
the good man is elect. I cannot forbear praising exceedingly the poet
of Agrigentum, who celebrates faith as follows:
“Friends, I know, then, that there is truth in the myths
Which I will relate. But very difficult to men,
And irksome to the mind, is the attempt of faith.”[716]
Wherefore also the apostle exhorts, “that your faith should not be in
the wisdom of men,” who profess to persuade, “but in the power of
God,”[717] which alone without proofs, by mere faith, is able to save.
“For the most approved of those that are reputable knows how to
keep watch. And justice will apprehend the forgers and witnesses of
lies,” says the Ephesian.[718] For he, having derived his knowledge
from the barbarian philosophy, is acquainted with the purification by
fire of those who have led bad lives, which the Stoics afterwards
called the Conflagration (ἐκπύρωσις), in which also they teach that
each will arise exactly as he was, so treating of the resurrection;
while Plato says as follows, that the earth at certain periods is
purified by fire and water: “There have been many destructions of
men in many ways; and there shall be very great ones by fire and
water; and others briefer by innumerable causes.” And after a little
he adds: “And, in truth, there is a change of the objects which
revolve about earth and heaven; and in the course of long periods
there is the destruction of the objects on earth by a great
conflagration.” Then he subjoins respecting the deluge: “But when,
again, the gods deluge the earth to purify it with water, those on the
mountains, herdsmen and shepherds, are saved; those in your cities
are carried down by the rivers into the sea.” And we showed in the
first Miscellany that the philosophers of the Greeks are called
thieves, inasmuch as they have taken without acknowledgment their
principal dogmas from Moses and the prophets. To which also we
shall add, that the angels who had obtained the superior rank,
having sunk into pleasures, told to the women the secrets which had
come to their knowledge; while the rest of the angels concealed
them, or rather, kept them against the coming of the Lord. Thence
emanated the doctrine of providence, and the revelation of high
things; and prophecy having already been imparted to the
philosophers of the Greeks, the treatment of dogma arose among
the philosophers, sometimes true when they hit the mark, and
sometimes erroneous, when they comprehended not the secret of
the prophetic allegory. And this it is proposed briefly to indicate in
running over the points requiring mention. Faith, then, we say, we
are to show must not be inert and alone, but accompanied with
investigation. For I do not say that we are not to inquire at all. For
“Search, and thou shalt find,”[719] it is said.
“What is sought may be captured,
But what is neglected escapes,”
according to Sophocles.
The like also says Menander the comic poet:
“All things sought,
The wisest say, need anxious thought.”
But we ought to direct the visual faculty of the soul aright to
discovery, and to clear away obstacles; and to cast clean away
contention, and envy, and strife, destined to perish miserably from
among men.
For very beautifully does Timon of Phlius write:
“And Strife, the Plague of Mortals, stalks vainly shrieking,
The sister of Murderous Quarrel and Discord,
Which rolls blindly over all things. But then
It sets its head towards men, and casts them on hope.”
Then a little below he adds:
“For who hath set these to fight in deadly strife?
A rabble keeping pace with Echo; for, enraged at those silent,
It raised an evil disease against men, and many perished;”
of the speech which denies what is false, and of the dilemma, of
that which is concealed, of the Sorites, and of the Crocodilean, of
that which is open, and of ambiguities and sophisms. To inquire,
then, respecting God, if it tend not to strife, but to discovery, is
salutary. For it is written in David, “The poor eat, and shall be filled;
and they shall praise the Lord that seek Him. Your heart shall live for
ever.”[720] For they who seek Him after the true search, praising the
Lord, shall be filled with the gift that comes from God, that is,
knowledge. And their soul shall live; for the soul is figuratively
termed the heart, which ministers life: for by the Son is the Father
known.
We ought not to surrender our ears to all who speak and write
rashly. For cups also, which are taken hold of by many by the ears,
are dirtied, and lose the ears; and besides, when they fall they are
broken. In the same way also, those, who have polluted the pure
hearing of faith by many trifles, at last becoming deaf to the truth,
become useless and fall to the earth. It is not, then, without reason
that we commanded boys to kiss their relations, holding them by the
ears; indicating this, that the feeling of love is engendered by
hearing. And “God,” who is known to those who love, “is love,”[721]
as “God,” who by instruction is communicated to the faithful, “is
faithful;”[722] and we must be allied to Him by divine love: so that by
like we may see like, hearing the word of truth guilelessly and
purely, as children who obey us. And this was what he, whoever he
was, indicated who wrote on the entrance to the temple at
Epidaurus the inscription:
“Pure he must be who goes within
The incense-perfumed fane.”
And purity is “to think holy thoughts.” “Except ye become as these
little children, ye shall not enter,” it is said, “into the kingdom of
heaven.”[723] For there the temple of God is seen established on
three foundations—faith, hope, and love.
CHAPTER II.
ON HOPE.
Respecting faith we have adduced sufficient testimonies of
writings among the Greeks. But in order not to exceed bounds,
through eagerness to collect a very great many also respecting hope
and love, suffice it merely to say that in the Crito Socrates, who
prefers a good life and death to life itself, thinks that we have hope
of another life after death.
Also in the Phædrus he says, “That only when in a separate state
can the soul become partaker of the wisdom which is true, and
surpasses human power; and when, having reached the end of hope
by philosophic love, desire shall waft it to heaven, then,” says he,
“does it receive the commencement of another, an immortal life.”
And in the Symposium he says, “That there is instilled into all the
natural love of generating what is like, and in men of generating
men alone, and in the good man of the generation of the
counterpart of himself. But it is impossible for the good man to do
this without possessing the perfect virtues, in which he will train the
youth who have recourse to him.” And as he says in the Theætetus,
“He will beget and finish men. For some procreate by the body,
others by the soul;” since also with the barbarian philosophers to
teach and enlighten is called to regenerate; and “I have begotten
you in Jesus Christ,”[724] says the good apostle somewhere.
Empedocles, too, enumerates friendship among the elements,
conceiving it as a combining love:
“Which do you look at with your mind; and don’t sit gaping with your eyes.”
Parmenides, too, in his poem, alluding to hope, speaks thus:
“Yet look with the mind certainly on what is absent as present,
For it will not sever that which is from the grasp it has of that which is
Not, even if scattered in every direction over the world or combined.”
CHAPTER III.
THE OBJECTS OF FAITH AND HOPE PERCEIVED BY THE MIND ALONE.
For he who hopes, as he who believes, sees intellectual objects
and future things with the mind. If, then, we affirm that aught is
just, and affirm it to be good, and we also say that truth is
something, yet we have never seen any of such objects with our
eyes, but with our mind alone. Now the Word of God says, “I am the
truth.”[725] The Word is then to be contemplated by the mind. “Do
you aver,” it was said,[726] “that there are any true philosophers?”
“Yes,” said I, “those who love to contemplate the truth.” In the
Phædrus also, Plato, speaking of the truth, shows it as an idea. Now
an idea is a conception of God; and this the barbarians have termed
the Word of God. The words are as follow: “For one must then dare
to speak the truth, especially in speaking of the truth. For the
essence of the soul, being colourless, formless, and intangible, is
visible only to God,[727] its guide.” Now the Word issuing forth was
the cause of creation; then also he generated himself, “when the
Word had become flesh,”[728] that He might be seen. The righteous
man will seek the discovery that flows from love, to which if he
hastes he prospers. For it is said, “To him that knocketh, it shall be
opened: ask, and it shall be given to you.”[729] “For the violent that
storm the kingdom”[730] are not so in disputatious speeches; but by
continuance in a right life and unceasing prayers, are said “to take it
by force,” wiping away the blots left by their previous sins.
“You may obtain wickedness, even in great abundance.[731] And
him who toils God helps; For the gifts of the Muses, hard to win, Lie
not before you, for any one to bear away.”
The knowledge of ignorance is, then, the first lesson in walking
according to the Word. An ignorant man has sought, and having
sought, he finds the teacher; and finding has believed, and believing
has hoped; and henceforward having loved, is assimilated to what
was loved—endeavouring to be what he first loved. Such is the
method Socrates shows Alcibiades, who thus questions: “Do you not
think that I shall know about what is right otherwise?” “Yes, if you
have found out.” “But you don’t think I have found out?” “Certainly,
if you have sought.” “Then you don’t think that I have sought?” “Yes,
if you think you do not know.”[732] So with the lamps of the wise
virgins, lighted at night in the great darkness of ignorance, which the
Scripture signified by “night.” Wise souls, pure as virgins,
understanding themselves to be situated amidst the ignorance of the
world, kindle the light, and rouse the mind, and illumine the
darkness, and dispel ignorance, and seek truth, and await the
appearance of the Teacher.
“The mob, then,” said I, “cannot become a philosopher.”[733]
“Many rod-bearers there are, but few Bacchi,” according to Plato.
“For many are called, but few chosen.”[734] “Knowledge is not in
all,”[735] says the apostle. “And pray that we may be delivered from
unreasonable and wicked men: for all men have not faith.”[736] And
the Poetics of Cleanthes, the Stoic, writes to the following effect:
“Look not to glory, wishing to be suddenly wise,
And fear not the undiscerning and rash opinion of the many;
For the multitude has not an intelligent, or wise, or right judgment,
And it is in few men that you will find this.”[737]
And more sententiously the comic poet briefly says:
“It is a shame to judge of what is right by much noise.”
For they heard, I think, that excellent wisdom, which says to us,
“Watch your opportunity in the midst of the foolish, and in the midst
of the intelligent continue.”[738] And again, “The wise will conceal
sense.”[739] For the many demand demonstration as a pledge of
truth, not satisfied with the bare salvation by faith.
“But it is strongly incumbent to disbelieve the dominant wicked,
And as is enjoined by the assurance of our muse,
Know by dissecting the utterance within your breast.”
“For this is habitual to the wicked,” says Empedocles, “to wish to
overbear what is true by disbelieving it.” And that our tenets are
probable and worthy of belief, the Greeks shall know, the point being
more thoroughly investigated in what follows. For we are taught
what is like by what is like. For says Solomon, “Answer a fool
according to his folly.”[740] Wherefore also, to those that ask the
wisdom that is with us, we are to hold out things suitable, that with
the greatest possible ease they may, through their own ideas, be
likely to arrive at faith in the truth. For “I became all things to all
men, that I might gain all men.”[741] Since also “the rain” of the
divine grace is sent down “on the just and the unjust.”[742] “Is He
the God of the Jews only, and not also of the Gentiles? Yes, also of
the Gentiles: if indeed He is one God,”[743] exclaims the noble
apostle.
CHAPTER IV.
DIVINE THINGS WRAPPED UP IN FIGURES BOTH IN THE SACRED AND IN
HEATHEN WRITERS.
But since they will believe neither in what is good justly nor in
knowledge unto salvation, we ourselves reckoning what they claim
as belonging to us, because all things are God’s; and especially since
what is good proceeded from us to the Greeks, let us handle those
things as they are capable of hearing. For intelligence or rectitude
this great crowd estimates not by truth, but by what they are
delighted with. And they will be pleased not more with other things
than with what is like themselves. For he who is still blind and dumb,
not having understanding, or the undazzled and keen vision of the
contemplative soul, which the Saviour confers, like the uninitiated at
the mysteries, or the unmusical at dances, not being yet pure and
worthy of the pure truth, but still discordant and disordered and
material, must stand outside of the divine choir. “For we compare
spiritual things with spiritual.”[744] Wherefore, in accordance with the
method of concealment, the truly sacred Word, truly divine and most
necessary for us, deposited in the shrine of truth, was by the
Egyptians indicated by what were called among them adyta, and by
the Hebrews by the veil. Only the consecrated—that is, those
devoted to God, circumcised in the desires of the passions for the
sake of love to that which is alone divine—were allowed access to
them. For Plato also thought it not lawful for “the impure to touch
the pure.”
Thence the prophecies and oracles are spoken in enigmas, and
the mysteries are not exhibited incontinently to all and sundry, but
only after certain purifications and previous instructions.
“For the Muse was not then
Greedy of gain or mercenary;
Nor were Terpsichore’s sweet,
Honey-toned, silvery soft-voiced
Strains made merchandise of.”
Now those instructed among the Egyptians learned first of all that
style of the Egyptian letters which is called Epistolographic; and
second, the Hieratic, which the sacred scribes practise; and finally,
and last of all, the Hieroglyphic, of which one kind which is by the
first elements is literal (Kyriologic), and the other Symbolic. Of the
Symbolic, one kind speaks literally by imitation, and another writes
as it were figuratively; and another is quite allegorical, using certain
enigmas.
Wishing to express Sun in writing, they make a circle; and Moon,
a figure like the Moon, like its proper shape. But in using the
figurative style, by transposing and transferring, by changing and by
transforming in many ways as suits them, they draw characters. In
relating the praises of the kings in theological myths, they write in
anaglyphs.[745] Let the following stand as a specimen of the third
species—the Enigmatic. For the rest of the stars, on account of their
oblique course, they have figured like the bodies of serpents; but the
sun like that of a beetle, because it makes a round figure of ox-
dung, and rolls it before its face. And they say that this creature lives
six months under ground, and the other division of the year above
ground, and emits its seed into the ball, and brings forth; and that
there is not a female beetle. All then, in a word, who have spoken of
divine things, both Barbarians and Greeks, have veiled the first
principles of things, and delivered the truth in enigmas, and
symbols, and allegories, and metaphors, and such like tropes. Such
also are the oracles among the Greeks. And the Pythian Apollo is
called Loxias. Also the maxims of those among the Greeks called
wise men, in a few sayings indicate the unfolding of matter of
considerable importance. Such certainly is that maxim, “Spare
Time:” either because life is short, and we ought not to expend this
time in vain; or, on the other hand, it bids you spare your personal
expenses; so that, though you live many years, necessaries may not
fail you. Similarly also the maxim “Know thyself” shows many things;
both that thou art mortal, and that thou wast born a human being;
and also that, in comparison with the other excellences of life, thou
art of no account, because thou sayest that thou art rich or
renowned; or, on the other hand, that, being rich or renowned, you
are not honoured on account of your advantages alone. And it says,
Know for what thou wert born, and whose image thou art; and what
is thy essence, and what thy creation, and what thy relation to God,
and the like. And the Spirit says by Isaiah the prophet, “I will give
thee treasures, hidden, dark.”[746] Now wisdom, hard to hunt, is the
treasures of God and unfailing riches. But those, taught in theology
by those prophets, the poets, philosophize much by way of a hidden
sense. I mean Orpheus, Linus, Musæus, Homer, and Hesiod, and
those in this fashion wise. The persuasive style of poetry is for them
a veil for the many. Dreams and signs are all more or less obscure to
men, not from jealousy (for it were wrong to conceive of God as
subject to passions), but in order that research, introducing to the
understanding of enigmas, may haste to the discovery of truth. Thus
Sophocles the tragic poet somewhere says:
“And God I know to be such an one,
Ever the revealer of enigmas to the wise,
But to the perverse bad, although a teacher in few words,”—
putting bad instead of simple. Expressly then respecting all our
Scripture, as if spoken in a parable, it is written in the Psalms, “Hear,
O my people, my law: incline your ear to the words of my mouth. I
will open my mouth in parables, I will utter my problems from the
beginning.”[747] Similarly speaks the noble apostle to the following
effect: “Howbeit we speak wisdom among those that are perfect; yet
not the wisdom of this world, nor of the princes of this world, that
come to nought. But we speak the wisdom of God hidden in a
mystery; which none of the princes of this world knew. For had they
known it, they would not have crucified the Lord of glory.”[748]
The philosophers did not exert themselves in contemning the
appearance of the Lord. It therefore follows that it is the opinion of
the wise among the Jews which the apostle inveighs against it.
Wherefore he adds, “But we preach, as it is written, what eye hath
not seen, and ear hath not heard, and hath not entered into the
heart of man, what God hath prepared for them that love Him. For
God hath revealed it to us by the Spirit. For the Spirit searcheth all
things, even the deep things of God.”[749] For he recognises the
spiritual man and the gnostic as the disciple of the Holy Spirit
dispensed by God, which is the mind of Christ. “But the natural man
receiveth not the things of the Spirit, for they are foolishness to
him.”[750] Now the apostle, in contradistinction to gnostic perfection,
calls the common faith the foundation, and sometimes milk, writing
on this wise: “Brethren, I could not speak to you as to spiritual, but
as to carnal, to babes in Christ. I have fed you with milk, not with
meat: for ye were not able. Neither yet are ye now able. For ye are
yet carnal: for whereas there is among you envy and strife, are ye
not carnal, and walk as men?”[751] Which things are the choice of
those men who are sinners. But those who abstain from these things
give their thoughts to divine things, and partake of gnostic food.
“According to the grace,” it is said, “given to me as a wise master
builder, I have laid the foundation. And another buildeth on it gold
and silver, precious stones.”[752] Such is the gnostic superstructure
on the foundation of faith in Christ Jesus. But “the stubble, and the
wood, and the hay,” are the additions of heresies. “But the fire shall
try every man’s work, of what sort it is.” In allusion to the gnostic
edifice also in the Epistle to the Romans, he says, “For I desire to
see you, that I may impart unto you a spiritual gift, that ye may be
established.”[753] It was impossible that gifts of this sort could be
written without disguise.
CHAPTER V.
ON THE SYMBOLS OF PYTHAGORAS.
Now the Pythagorean symbols were connected with the Barbarian
philosophy in the most recondite way. For instance, the Samian
counsels “not to have a swallow in the house;” that is, not to receive
a loquacious, whispering, garrulous man, who cannot contain what
has been communicated to him. “For the swallow, and the turtle,
and the sparrows of the field, know the times of their entrance,”[754]
says the Scripture; and one ought never to dwell with trifles. And the
turtle-dove murmuring shows the thankless slander of fault-finding,
and is rightly expelled the house.
“Don’t mutter against me, sitting by one in one place, another in
another.”[755]
The swallow too, which suggests the fable of Pandion, seeing it is
right to detest the incidents reported of it, some of which we hear
Tereus suffered, and some of which he inflicted. It pursues also the
musical grasshoppers, whence he who is a persecutor of the word
ought to be driven away.
“By sceptre-bearing Here, whose eye surveys Olympus,
I have a trusty closet for tongues,”
says Poetry. Æschylus also says:
“But I, too, have a key as a guard on my tongue.”
Again Pythagoras commanded, “When the pot is lifted off the fire,
not to leave its mark in the ashes, but to scatter them;” and “people
on getting up from bed, to shake the bed-clothes.” For he intimated
that it was necessary not only to efface the mark, but not to leave
even a trace of anger; and that on its ceasing to boil, it was to be
composed, and all memory of injury to be wiped out. “And let not
the sun,” says the Scripture, “go down upon your wrath.”[756] And
he that said, “Thou shalt not desire,”[757] took away all memory of
wrong; for wrath is found to be the impulse of concupiscence in a
mild soul, especially seeking irrational revenge. In the same way
“the bed is ordered to be shaken up,” so that there may be no
recollection of effusion in sleep, or sleep in the day-time; nor,
besides, of pleasure during the night. And he intimated that the
vision of the dark ought to be dissipated speedily by the light of
truth. “Be angry, and sin not,” says David, teaching us that we ought
not to assent to the impression, and not to follow it up by action,
and so confirm wrath.
Again, “Don’t sail on land” is a Pythagorean saying, and shows
that taxes and similar contracts, being troublesome and fluctuating,
ought to be declined. Wherefore also the Word says that the tax-
gatherers shall be saved with difficulty.[758]
And again, “Don’t wear a ring, nor engrave on it the images of
the gods,” enjoins Pythagoras; as Moses ages before enacted
expressly, that neither a graven, nor molten, nor moulded, nor
painted likeness should be made; so that we may not cleave to
things of sense, but pass to intellectual objects: for familiarity with
the sight disparages the reverence of what is divine; and to worship
that which is immaterial by matter, is to dishonour it by sense.
Wherefore the wisest of the Egyptian priests decided that the temple
of Athene should be hypæthral, just as the Hebrews constructed the
temple without an image. And some, in worshipping God, make a
representation of heaven containing the stars; and so worship,
although Scripture says, “Let us make man in our image and
likeness.”[759] I think it worth while also to adduce the utterance of
Eurysus the Pythagorean, which is as follows, who in his book On
Fortune, having said that the “Creator, on making man, took Himself
as an exemplar,” added, “And the body is like the other things, as
being made of the same material, and fashioned by the best
workman, who wrought it, taking Himself as the archetype.” And, in
fine, Pythagoras and his followers, with Plato also, and most of the
other philosophers, were best acquainted with the Lawgiver, as may
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    The Princeton Guideto Evolution
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    princeton university press Princeton& Oxford The Princeton Guide to Evolution editor in chief Jonathan B. Losos Harvard University editors David A. Baum University of Wisconsin, Madison Douglas J. Futuyma Stony Brook University Hopi E. Hoekstra Harvard University Richard E. Lenski Michigan State University Allen J. Moore University of Georgia Catherine L. Peichel Fred Hutchinson Cancer 3esearch Center Dolph Schluter University of British Columbia Michael J. Whitlock University of British Columbia advisors Michael J. Donoghue Simon A. Levin Trudy F. C. Mackay Loren Rieseberg Joseph Travis Gregory A. Wray
  • 9.
    Copyright # 2014by Princeton University Press Published by Princeton University Press, 41 William Street, Princeton, New Jersey 08540 In the United Kingdom: Princeton University Press, 6 Oxford Street, Woodstock, Oxfordshire OX20 1TW press.princeton.edu All Rights Reserved Library of Congress Cataloging-in-Publication Data The Princeton guide to evolution / Jonathan B. Losos, Harvard University, editor in chief ; David A. Baum, University of Wisconsin, Madison, Douglas J. Futuyma, Stony Brook University, Hopi E. Hoekstra, Harvard University, Richard E. Lenski, Michigan State University, Allen J. Moore, University of Georgia, Catherine L. Peichel, Hutchinson Cancer Institute, Seattle, Dolph Schluter, University of British Columbia, Michael C. Whitlock, University of British Columbia, editors. pages cm Includes bibliographical references and index. ISBN 978-0-691-14977-6 (hardcover) 1. Evolution (Biology) I. Losos, Jonathan B. QH367.P85 2014 576.8—dc23 2013022360 British Library Cataloging-in-Publication Data is available This book has been composed in Sabon and Din Printed on acid-free paper. Printed in the United States of America 10 9 8 7 6 5 4 3 2 1
  • 10.
    Contents Preface vii Contributors ix SectionI Introduction 1 I.1 What Is Evolution? 3 I.2 The History of Evolutionary Thought 10 I.3 The Evidence for Evolution 28 I.4 From DNA to Phenotypes 40 Section II Phylogenetics and the History of Life 47 II.1 Interpretation of Phylogenetic Trees 51 II.2 Phylogenetic Inference 60 II.3 Molecular Clock Dating 67 II.4 Historical Biogeography 75 II.5 Phylogeography 82 II.6 Concepts in Character Macroevolution: Adaptation, Homology, and Evolvability 89 II.7 Using Phylogenies to Study Phenotypic Evolution: Comparative Methods and Tests of Adaptation 100 II.8 Taxonomy in a Phylogenetic Framework 106 II.9 The Fossil Record 112 II.10 The Origin of Life 120 II.11 Evolution in the Prokaryotic Grade 127 II.12 Origin and Diversification of Eukaryotes 136 II.13 Major Events in the Evolution of Land Plants 143 II.14 Major Events in the Evolution of Fungi 152 II.15 Origin and Early Evolution of Animals 159 II.16 Major Events in the Evolution of Arthropods 167 II.17 Major Features of Tetrapod Evolution 174 II.18 Human Evolution 183 Section III Natural Selection and Adaptation 189 III.1 Natural Selection, Adaptation, and Fitness: Overview 193 III.2 Units and Levels of Selection 200 III.3 Theory of Selection in Populations 206 III.4 Kin Selection and Inclusive Fitness 215 III.5 Phenotypic Selection on Quantitative Traits 221 III.6 Responses to Selection: Experimental Populations 230 III.7 Responses to Selection: Natural Populations 238 III.8 Evolutionary Limits and Constraints 247 III.9 Evolution of Modifier Genes and Biological Systems 253 III.10 Evolution of Reaction Norms 261 III.11 Evolution of Life Histories 268 III.12 Evolution of Form and Function 276 III.13 Biochemical and Physiological Adaptations 282 III.14 Evolution of the Ecological Niche 288 III.15 Adaptation to the Biotic Environment 298 Section IV Evolutionary Processes 305 IV.1 Genetic Drift 307 IV.2 Mutation 315 IV.3 Geographic Variation, Population Structure, and Migration 321 IV.4 Recombination and Sex 328 IV.5 Genetic Load 334 IV.6 Inbreeding 340 IV.7 Selfish Genetic Elements and Genetic Conflict 347 IV.8 Evolution of Mating Systems: Outcrossing versus Selfing 356 Section V Genes, Genomes, Phenotypes 363 V.1 Molecular Evolution 367 V.2 Genome Evolution 374 V.3 Comparative Genomics 380 V.4 Evolution of Sex Chromosomes 387 V.5 Gene Duplication 397 V.6 Evolution of New Genes 406 V.7 Evolution of Gene Expression 413 V.8 Epigenetics 420
  • 11.
    V.9 Evolution ofMolecular Networks 428 V.10 Evolution and Development: Organisms 436 V.11 Evolution and Development: Molecules 444 V.12 Genetics of Phenotypic Evolution 452 V.13 Dissection of Complex Trait Evolution 458 V.14 Searching for Adaptation in the Genome 466 V.15 Ancient DNA 475 Section VI Speciation and Macroevolution 483 VI.1 Species and Speciation 489 VI.2 Speciation Patterns 496 VI.3 Geography, Range Evolution, and Speciation 504 VI.4 Speciation and Natural Selection 512 VI.5 Speciation and Sexual Selection 520 VI.6 Gene Flow, Hybridization, and Speciation 529 VI.7 Coevolution and Speciation 535 VI.8 Genetics of Speciation 543 VI.9 Speciation and Genome Evolution 549 VI.10 Adaptive Radiation 559 VI.11 Macroevolutionary Rates 567 VI.12 Macroevolutionary Trends 573 VI.13 Causes and Consequences of Extinction 579 V1.14 Species Selection 586 VI.15 Key Evolutionary Innovations 592 VI.16 Evolution of Communities 599 Section VII Evolution of Behavior, Society, and Humans 605 VII.1 Genes, Brains, and Behavior 609 VII.2 Evolution of Hormones and Behavior 616 VII.3 Game Theory and Behavior 624 VII.4 Sexual Selection and Its Impact on Mating Systems 632 VII.5 Sexual Selection: Male-Male Competition 641 VII.6 Sexual Selection: Mate Choice 647 VII.7 Evolution of Communication 655 VII.8 Evolution of Parental Care 663 VII.9 Cooperation and Conflict: Microbes to Humans 671 VII.10 Cooperative Breeding 677 VII.11 Human Behavioral Ecology 683 VII.12 Evolutionary Psychology 690 VII.13 Evolution of Eusociality 697 VII.14 Cognition: Phylogeny, Adaptation, and By-Products 703 VII.15 Evolution of Apparently Nonadaptive Behavior 710 VII.16 Aging and Menopause 718 Section VIII Evolution and Modern Society 727 VIII.1 Evolutionary Medicine 733 VIII.2 Evolution of Parasite Virulence 741 VIII.3 Evolution of Antibiotic Resistance 747 VIII.4 Evolution and Microbial Forensics 754 VIII.5 Domestication and the Evolution of Agriculture 760 VIII.6 Evolution and Conservation 766 VIII.7 Directed Evolution 774 VIII.8 Evolution and Computing 780 VIII.9 Linguistics and the Evolution of Human Language 786 VIII.10 Cultural Evolution 795 VIII.11 Evolution and Notions of Human Race 801 VIII.12 The Future of Human Evolution 809 VIII.13 Evolution and Religion 817 VIII.14 Creationism and Intelligent Design 825 VIII.15 Evolution and the Media 832 Index 837 vi Contents
  • 12.
    Preface For more than150 years, since the publication of On the Origin of Species, biologists have focused on under- standing the evolutionary chronicle of diversification and extinction, and the underlying evolutionary processes that have produced it. Although progress in evolutionary biology has been steady since Darwin’s time, develop- ments in the last 20 years have ushered in a golden era of evolutionary study in which biologists are on the brink of answeringmanyofthefundamentalquestionsinthefield. These advances have come from a confluence of tech- nological and conceptual innovations. In the laboratory, the rapid and inexpensive sequencing of large amounts of DNA is producing a wealth of data on the genomes of many species; comparisons of these genomes are al- lowing scientists to pinpoint the specific genetic changes that have occurred over the course of evolution. In par- allel, spectacular fossil discoveries have filled many of the most critical gaps in our documentation of the evo- lutionary pageant, detailing how whales evolved from land-living animals, snakes from their four-legged lizard forebears, and humans from our primate ancestors. In addition, providing the data that Darwin could only imagine, field biologists are now tracking populations, directly documenting natural selection as it occurs, and monitoring the resulting evolutionary changes that oc- cur from one generation to the next. At the same time, evolutionary biology is making an impactthroughouthumansociety.Manycurrentissues— such as the rise of new diseases, the increasedresistance of pests and microorganisms to efforts to control them, and the effect of changing environmental conditions on nat- ural populations—revolve around aspects of natural se- lection and evolutionary change. Many disparate areas of modern life—medicine, the legal system, computing— increasingly employ evolutionary thinking and use meth- odsdevelopedinevolutionarybiology.Paradoxically,even as our understanding of evolution and its importance to society has never been greater, substantial proportions of thepopulationinanumberofcountries—mostnotablythe UnitedStatesandTurkey—disputethescientificfindingsof evolutionary biologists and resist the teaching of evolution in schools. This volume follows on the success of The Princeton Guide to Ecology, edited by Simon Levin. Published in 2009, the ecology guide has proven valuable to a wide range of readers, from professional ecologists and graduate students to land planners, economists, and social scientists. With this model in mind, we set out to produce a guide that would be accessible and useful to students and scientists in evolutionary biology and re- lated disciplines, as well as to anyone with a serious in- terest in evolution. What makes this volume stand out is the breadth and depth of our 107 chapters, each written by authorities in their respective field. In addition, the articles balance accessibility with depth of analysis, making the Guide a valuable reference for a broad au- dience. Certainly, some articles are more technical than others, but readers can easily select chapters appropriate for their interests and expertise. The Guide is divided into eight sections. The intro- ductory section includes four chapters covering the basics of evolution: what it is, the history of its study, the evi- dence for its occurrence, and a basic primer of genetic and phenotypic variation. The following seven sections cover the major areas of evolutionary biology, each beginning with a synoptic overview by the section editor. Section II: PhylogeneticsandtheHistoryofLife,coversthehistoryof life andhowitisstudied.Itincludeschaptersontheevolu- tion of each of the major forms of life, as well as on the studyoflife’shistorythroughtheexaminationofthefossil record and the construction of phylogenetic trees that detail the relationships among species and higher taxa. Section III: Selection and Adaptation, moves to evolu- tionary processes, focusing on natural selection, the pre- sumed primary driver of evolutionary change. Section IV: Evolutionary Processes, covers gene flow, genetic drift, andnonrandommating.SectionV:Genes,Genomes,Phe- notypes, examines the link between genes and phenotypes and how they evolve, focusing on the rapid growth of knowledge and continuing research in genetics and de- velopmental biology and the relationships of these fields to evolutionary biology. Section VI: Speciation and Mac- roevolution, moves the focus to the species level and above,emphasizingtheoriginofspecies—thatis,speciation —andevolutionarychangethatdriveslarge-scalechanges in the history of life through time, such as the rise of particular taxa and the extinction of others. Section VII: Evolution of Behavior, Society, and Humans, focuses on behavioral and social interactions that occur within spe- cies, including competition for mating success (referredto
  • 13.
    as sexual selection)and the evolution of traits such as parental care, communication, and altruism. Although chapters in this section are broad in taxonomic scope, many have particular relevance to human biology. Fi- nally, Section VIII: Evolution and Modern Society, ad- dresses how evolutionary biology directly affects the health and welfare of humans today. This volume could not have been possible without the efforts of the editors and authors, whose work was instru- mental to such a wide-ranging and authoritative work. Developmentoftheguidealsobenefitedimmenselyfromthe wisdom of our advisors, Michael Donoghue, Simon Levin, Trudy Mackay, Loren Rieseberg, Joseph Travis, and Greg Wray.Inaddition,theeditorialstaffatPrincetonUniversity Press was indispensable. The entire project was skillfully overseenbyexecutiveeditorAnneSavarese,andday-to-day management moved smoothly and efficiently under the watchful eyes of editorial assistants Diana Goovaerts and Sarah David, and production editor Karen Carter. Wemournthelossandgratefullyacknowledgethecon- tribution of our distinguished colleague Farish Jenkins, who died on November 11, 2012. Jonathan B. Losos viii Preface
  • 14.
    Contributors Aneil F. Agrawal,Department of Ecology and Evolutionary Biology, University of Toronto IV.5 GENETIC LOAD Michael E. Alfaro, Department of Ecology and Evolutionary Biology, University of California, Los Angeles VI.15 KEY EVOLUTIONARY INNOVATIONS Garland E. Allen, Department of Biology, Washington University in St. Louis I.2 THE HISTORY OF EVOLUTIONARY THOUGHT Dan I. Andersson, Department of Medical Biochemistry and Microbiology, Uppsala University VIII.3 EVOLUTION OF ANTIBIOTIC RESISTANCE Michael J. Angilletta Jr., School of Life Sciences, Arizona State University III.13 BIOCHEMICAL AND PHYSIOLOGICAL ADAPTATIONS Charles F. Aquadro, Department of Molecular Biology and Genetics, Cornell University V.1 MOLECULAR EVOLUTION Jonathan W. Atwell, Department of Biology, Indiana University VII.2 EVOLUTION OF HORMONES AND BEHAVIOR Francisco J. Ayala, Department of Ecology and Evolutionary Biology, University of California, Irvine VIII.13 EVOLUTION AND RELIGION Doris Bachtrog, Department of Integrative Biology, University of California, Berkeley V.4 EVOLUTION OF SEX CHROMOSOMES Charles F. Baer, Department of Biology, University of Florida IV.2 MUTATION Nathan W. Bailey, School of Biology, University of St. Andrews VII.15 EVOLUTION OF APPARENTLY NONADAPTIVE BEHAVIOR Timothy G. Barraclough, Division of Ecology and Evolution, Imperial College London VI.2 SPECIATION PATTERNS Spencer C. H. Barrett, Department of Ecology and Evolutionary Biology, University of Toronto IV.8 EVOLUTION OF MATING SYSTEMS: OUTCROSSING VERSUS SELFING N. H. Barton, Institute of Science and Technology Austria IV.4 RECOMBINATION AND SEX David A. Baum, Department of Botany, University of Wisconsin, Madison II PHYLOGENETICS AND THE HISTORY OF LIFE Graham Bell, Department of Biology, McGill University III.6 RESPONSES TO SELECTION: EXPERIMENTAL POPULATIONS Yehuda Ben-Shahar, Department of Biology, Washington University in St. Louis VII.1 GENES, BRAINS, AND BEHAVIOR Michael J. Benton, School of Earth Sciences, University of Bristol VI.13 CAUSES AND CONSEQUENCES OF EXTINCTION Janette W. Boughman, Department of Zoology, Michigan State University VI.5 SPECIATION AND SEXUAL SELECTION Paul M. Brakefield, Department of Zoology, University of Cambridge V.10 EVOLUTION AND DEVELOPMENT: ORGANISMS Edmund D. Brodie III, Department of Biology, University of Virginia III.5 PHENOTYPIC SELECTION ON QUANTITATIVE TRAITS C. Alex Buerkle, Department of Botany and Program in Ecology, University of Wyoming VI.6 GENE FLOW, HYBRIDIZATION, AND SPECIATION Michael A. Cant, Biosciences, University of Exeter VII.10 COOPERATIVE BREEDING Paulyn Cartwright, Department of Ecology and Evolutionary Biology, University of Kansas II.15 ORIGIN AND EARLY EVOLUTION OF ANIMALS Amy Cavanaugh, Department of Biological Sciences, University of Wisconsin, Rock County VIII.5 DOMESTICATION AND THE EVOLUTION OF AGRICULTURE Michel Chapuisat, Department of Ecology and Evolution, University of Lausanne VII.13 EVOLUTION OF EUSOCIALITY Deborah Charlesworth, School of Biological Sciences, University of Edinburgh IV.6 INBREEDING Julia Clarke, Jackson School of Geosciences, University of Texas at Austin II.8 TAXONOMY IN A PHYLOGENETIC FRAMEWORK Peter R. Crane, School of Forestry and Environmental Studies, Yale University II.13 MAJOR EVENTS IN THE EVOLUTION OF LAND PLANTS
  • 15.
    Cameron R. Currie,Department of Bacteriology, University of Wisconsin, Madison VIII.5 DOMESTICATION AND THE EVOLUTION OF AGRICULTURE David Deamer, Department of Biomolecular Engineering, University of California, Santa Cruz II.10 THE ORIGIN OF LIFE Michael J. Donoghue, Department of Ecology and Evolutionary Biology, Yale University II.4 HISTORICAL BIOGEOGRAPHY Dieter Ebert, Zoological Institute, Universität Basel VIII.2 EVOLUTION OF PARASITE VIRULENCE Scott P. Egan, Department of Biological Sciences, University of Notre Dame VI.9 SPECIATION AND GENOME EVOLUTION Andrew D. Ellington, Department of Chemistry and Biochemistry, University of Texas at Austin VIII.7 DIRECTED EVOLUTION Jeffrey Feder, Department of Biological Sciences, University of Notre Dame VI.9 SPECIATION AND GENOME EVOLUTION Lila Fishman, Division of Biological Sciences, University of Montana IV.7 SELFISH GENETIC ELEMENTS AND GENETIC CONFLICT Douglas J. Futuyma, Department of Ecology and Evolution, Stony Brook University III NATURAL SELECTION AND ADAPTATION Dana H. Geary, Department of Geoscience, University of Wisconsin, Madison II.9 THE FOSSIL RECORD J. Peter Gogarten, Department of Molecular and Cell Biology, University of Connecticut II.11 EVOLUTION IN THE PROKARYOTIC GRADE Emma E. Goldberg, Biological Sciences, University of Illinois, Chicago VI.14 SPECIES SELECTION Peter R. Grant, Department of Ecology and Evolutionary Biology, Princeton University VI.10 ADAPTIVE RADIATION Michael D. Greenfield, Institut de Recherche sur la Biologie de l’Insecte, Université de Tours VII.7 EVOLUTION OF COMMUNICATION Elizabeth Hannon, Department of History and Philosophy of Science, University of Cambridge VIII.10 CULTURAL EVOLUTION Sara J. Hanson, Department of Biology and Program in Genetics, University of Iowa V.2 GENOME EVOLUTION Luke J. Harmon, Department of Biological Sciences, University of Idaho VI.11 MACROEVOLUTIONARY RATES Richard G. Harrison, Department of Ecology and Evolutionary Biology, Cornell University VI.1 SPECIES AND SPECIATION Marc D. Hauser, Independent Scholar VII.14 COGNITION: PHYLOGENY, ADAPTATION, AND BY- PRODUCTS John Hawks, Department of Anthropology, University of Wisconsin, Madison II.18 HUMAN EVOLUTION Philip Hedrick, School of Life Sciences, Arizona State University IV.1 GENETIC DRIFT Noel A. Heim, Department of Geology and Evolutionary Science, Stanford University II.9 THE FOSSIL RECORD Michael E. Hellberg, Department of Biological Sciences, Louisiana State University II.5 PHYLOGEOGRAPHY David S. Hibbett, Department of Biology, Clark University II.14 MAJOR EVENTS IN THE EVOLUTION OF FUNGI Hopi E. Hoekstra, Department of Organismic and Evolutionary Biology, Harvard University V GENES, GENOMES, PHENOTYPES Ary Hoffmann, Department of Genetics and Zoology, University of Melbourne III.8 LIMITS AND CONSTRAINTS Mark Holder, Department of Ecology and Evolutionary Biology, University of Kansas II.2 PHYLOGENETIC INFERENCE Kent E. Holsinger, Department of Ecology and Evolutionary Biology, University of Connecticut III.3 THEORY OF SELECTION IN POPULATIONS Robert D. Holt, Department of Ecology, University of Florida III.14 EVOLUTION OF THE ECOLOGICAL NICHE Robin Hopkins, Department of Integrative Biology, University of Texas at Austin VI.4 SPECIATION AND NATURAL SELECTION Gene Hunt, Department of Paleobiology Smithsonian Institution, National Museum of Natural History VI.12 MACROEVOLUTIONARY TRENDS John Jaenike, Department of Biology, University of Rochester IV.7 SELFISH GENETIC ELEMENTS AND GENETIC CONFLICT Farish A. Jenkins Jr., Late Professor of Biology, Harvard University II.17 MAJOR FEATURES OF TETRAPOD EVOLUTION Michael D. Jennions, Research School of Biology, Australian National University VII.6 SEXUAL SELECTION: MATE CHOICE Laura A. Katz, Department of Biological Sciences, Smith College II.12 ORIGIN AND DIVERSIFICATION OF EUKARYOTES x Contributors
  • 16.
    Paul Keim, Departmentof Biology, Northern Arizona University VIII.4 EVOLUTION AND MICROBIAL FORENSICS Laurent Keller, Department of Ecology and Evolution, University of Lausanne VII.13 EVOLUTION OF EUSOCIALITY Ellen D. Ketterson, Department of Biology, Indiana University VII.2 EVOLUTION OF HORMONES AND BEHAVIOR Joel G. Kingsolver, Department of Biology, University of North Carolina, Chapel Hill III.7 RESPONSES TO SELECTION: NATURAL POPULATIONS Hanna Kokko, Research School of Biology, Australian National University VII.6 SEXUAL SELECTION: MATE CHOICE Mathias Kölliker, Department of Environmental Sciences, University of Basel VII.8 EVOLUTION OF PARENTAL CARE Allan Larson, Department of Biology, Washington University in St. Louis II.6 CONCEPTS IN CHARACTER MACROEVOLUTION: ADAPTATION, HOMOLOGY, AND EVOLVABILITY Richard E. Lenski, Departments of Microbiology & Molecular Genetics, Zoology, and Crop and Soil Sciences, Michigan State University VIII EVOLUTION AND MODERN SOCIETY Andrew B. Leslie, School of Forestry and Environmental Studies, Yale University II.13 MAJOR EVENTS IN THE EVOLUTION OF LAND PLANTS Tim Lewens, Department of History and Philosophy of Science, University of Cambridge VIII.10 CULTURAL EVOLUTION John M. Logsdon Jr., Department of Biology and Program in Genetics, University of Iowa V.2 GENOME EVOLUTION Manyuan Long, Department of Ecology and Evolution, University of Chicago V.6 EVOLUTION OF NEW GENES Jonathan B. Losos, Department of Organismic and Evolutionary Biology, Harvard University I.1 WHAT IS EVOLUTION? David B. Lowry, Department of Integrative Biology, University of Texas at Austin VI.4 SPECIATION AND NATURAL SELECTION Virpi Lummaa, Department of Animal and Plant Sciences, University of Sheffield VII.11 HUMAN BEHAVIORAL ECOLOGY Florian Maderspacher, Current Biology, Elsevier, Inc. V.8 EPIGENETICS Gregory C. Mayer, Department of Biological Sciences, University of WisconsinParkside I.3 THE EVIDENCE FOR EVOLUTION Joel W. McGlothlin, Department of Biological Sciences, Virginia Polytechnic Institute and State University VII.2 EVOLUTION OF HORMONES AND BEHAVIOR Daniel McNabney, Department of Biology, University of Rochester VI.8 GENETICS OF SPECIATION John M. McNamara, School of Mathematics, University of Bristol VII.3 GAME THEORY AND BEHAVIOR Mark A. McPeek, Department of Biological Sciences, Dartmouth College VI.16 EVOLUTION OF COMMUNITIES Christine W. Miller, Department of Entomology and Nematology, University of Florida VII.5 SEXUAL SELECTION: MALE-MALE COMPETITION Antónia Monteiro, Department of Ecology and Evolutionary Biology, Yale University V.11 EVOLUTION AND DEVELOPMENT: MOLECULES Jacob A. Moorad, Department of Biology, Duke University VII.16 AGING AND MENOPAUSE Allen J. Moore, Department of Genetics, University of Georgia VII EVOLUTION OF BEHAVIOR, SOCIETY, AND HUMANS Patrik Nosil, Department of Ecology and Evolutionary Biology, University of Colorado, Boulder VI.9 SPECIATION AND GENOME EVOLUTION Samir Okasha, Department of Philosophy, University of Bristol III.2 UNITS AND LEVELS OF SELECTION Lorraine Olendzenski, Department of Biology, St. Lawrence University II.11 EVOLUTION IN THE PROKARYOTIC GRADE Kevin E. Omland, Department of Biological Sciences, University of Maryland, Baltimore County II.1 INTERPRETATION OF PHYLOGENETIC TREES H. Allen Orr, Department of Biology, University of Rochester VI.8 GENETICS OF SPECIATION Sarah P. Otto, Department of Zoology, University of British Columbia III.9 EVOLUTION OF MODIFIER GENES AND BIOLOGICAL SYSTEMS Mark Pagel, School of Biological Sciences, University of Reading VIII.9 LINGUISTICS AND THE EVOLUTION OF HUMAN LANGUAGE Laura Wegener Parfrey, Department of Chemistry and Biochemistry, University of Colorado, Boulder II.12 ORIGIN AND DIVERSIFICATION OF EUKARYOTES Bret A. Payseur, Laboratory of Genetics, University of Wisconsin, Madison V.13 DISSECTION OF COMPLEX TRAIT EVOLUTION Contributors xi
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    Talima Pearson, Departmentof Biological Sciences, Northern Arizona University VIII.4 EVOLUTION AND MICROBIAL FORENSICS Catherine L. Peichel, Fred Hutchinson Cancer Research Center, Seattle V GENES, GENOMES, PHENOTYPES; V.12 GENETICS OF PHENOTYPIC EVOLUTION Robert T. Pennock, Lyman Briggs College and Departments of Philosophy and Computer Science Engineering, Michigan State University VIII.8 EVOLUTION AND COMPUTING Dmitri A. Petrov, Department of Biology, Stanford University V.14 SEARCHING FOR ADAPTATION IN THE GENOME David W. Pfennig, Department of Biology, University of North Carolina, Chapel Hill III.7 RESPONSES TO SELECTION: NATURAL POPULATIONS Albert Phillimore, Institute of Evolutionary Biology, University of Edinburgh VI.3 GEOGRAPHY, RANGE EVOLUTION, AND SPECIATION Daniel E. L. Promislow, Department of Pathology, University of Washington VII.16 AGING AND MENOPAUSE Erik Quandt, Department of Chemistry and Biochemistry, University of Texas at Austin VIII.7 DIRECTED EVOLUTION David C. Queller, Department of Biology, Washington University in St. Louis III.4 KIN SELECTION AND INCLUSIVE FITNESS; VII.9 COOPERATION AND CONFLICT: MICROBES TO HUMANS Bruce Rannala, Department of Evolution and Ecology, University of California, Davis II.3 MOLECULAR CLOCK DATING Richard Ree, Botany Department, Field Museum of Natural History II.7 USING PHYLOGENIES TO STUDY PHENOTYPIC EVOLUTION: COMPARATIVE METHODS AND TESTS OF ADAPTATION David Reznick, Department of Biology, University of California, Riverside III.11 EVOLUTION OF LIFE HISTORIES Robert C. Richardson, Department of Philosophy, University of Cincinnati VII.12 EVOLUTIONARY PSYCHOLOGY Ophélie Ronce, Institut des Sciences de l’Evolution, Université Montpellier 2, Centre National de la Recherche Scientifique IV.3 GEOGRAPHIC VARIATION, POPULATION STRUCTURE, AND MIGRATION Nick J. Royle, Department of Biosciences, University of Exeter VII.8 EVOLUTION OF PARENTAL CARE Dolph Schluter, Department of Zoology, University of British Columbia VI SPECIATION AND MACROEVOLUTION Eugenie C. Scott, National Center for Science Education, Inc. VIII.14 CREATIONISM AND INTELLIGENT DESIGN H. Bradley Shaffer, Department of Ecology and Evolutionary Biology, University of California, Los Angeles VIII.6 EVOLUTION AND CONSERVATION Beth Shapiro, Department of Ecology and Evolutionary Biology, University of California, Santa Cruz V.15 ANCIENT DNA Mark L. Siegal, Department of Biology, New York University V.9 EVOLUTION OF MOLECULAR NETWORKS Per T. Smiseth, Institute of Evolutionary Biology, University of Edinburgh VII.8 EVOLUTION OF PARENTAL CARE Rhonda R. Snook, Department of Animal and Plant Sciences, University of Sheffield VII.4 SEXUAL SELECTION AND ITS IMPACT ON MATING SYSTEMS Jason E. Stajich, Department of Plant Pathology and Microbiology, University of California, Riverside V.3 COMPARATIVE GENOMICS Stephen C. Stearns, Department of Ecology and Evolutionary Biology, Yale University III.1 NATURAL SELECTION, ADAPTATION, AND FITNESS: OVERVIEW; III.10 EVOLUTION OF REACTION NORMS Joan E. Strassmann, Department of Biology, Washington University in St. Louis III.4 KIN SELECTION AND INCLUSIVE FITNESS; VII.9 COOPERATION AND CONFLICT: MICROBES TO HUMANS Sharon Y. Strauss, Department of Ecology and Evolution, University of California, Davis III.15 ADAPTATION TO THE BIOTIC ENVIRONMENT Alan R. Templeton, Department of Biology, Washington University in St. Louis, Department of Evolutionary and Environmental Biology, University of Haifa VIII.11 EVOLUTION AND NOTIONS OF HUMAN RACE; VIII.12 THE FUTURE OF HUMAN EVOLUTION John N. Thompson, Department of Ecology and Evolutionary Biology, University of California, Santa Cruz VI.7 COEVOLUTION AND SPECIATION Michelle D. Trautwein, Biodiversity Laboratory, North Carolina Museum of Natural Sciences II.16 MAJOR EVENTS IN THE EVOLUTION OF ARTHROPODS Paul E. Turner, Department of Ecology and Evolutionary Biology, Yale University VIII.1 EVOLUTIONARY MEDICINE Peter C. Wainwright, Department of Evolution and Ecology, University of California, Davis III.12 EVOLUTION OF FORM AND FUNCTION xii Contributors
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    Michael C. Whitlock,Department of Zoology, University of British Columbia I.4 FROM DNA TO PHENOTYPES; IV EVOLUTIONARY PROCESSES Brian M. Wiegmann, Department of Entomology, North Carolina State University II.16 MAJOR EVENTS IN THE EVOLUTION OF ARTHROPODS Patricia J. Wittkopp, Department of Ecology and Evolutionary Biology, University of Michigan V.7 EVOLUTION OF GENE EXPRESSION Ziheng Yang, Department of Genetics, Evolution, and Environment, University College, London II.3 MOLECULAR CLOCK DATING Jianzhi Zhang, Department of Ecology and Evolutionary Biology, University of Michigan V.5 GENE DUPLICATION Carl Zimmer, Environmental Studies Program, Yale University VIII.15 EVOLUTION AND THE MEDIA Contributors xiii
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  • 22.
    I.1 What Is Evolution? JonathanLosos OUTLINE 1. What is evolution? 2. Evolution: Pattern versus process 3. Evolution: More than changes in the gene pool 4. In the light of evolution 5. Critiques and the evidence for evolution 6. The pace of evolution 7. Evolution, humans, and society Evolution refers to change through time as species be- come modified and diverge to produce multiple descen- dant species. Evolution and natural selection are often conflated, but evolution is the historical occurrence of change, and natural selection is one mechanism—in most cases the most important—that can cause it. Recent years have seen a flowering in the field of evolutionary biology, and much has been learned about the causes and consequences of evolution. The two main pillars of our knowledge of evolution come from knowledge of the historicalrecordofevolutionarychange,deduceddirectly from the fossil record and inferred from examination of phylogeny, and from study of the process of evolutionary change, particularly the effect of natural selection. It is now apparent that whenselection is strong,evolution can proceed considerably more rapidly than was generally envisioned by Darwin. As a result, scientists are realizing that it is possible to conduct evolutionary experiments in real time. Recent developments in many areas, including molecular and developmental biology, have greatly ex- panded our knowledge and reaffirmed evolution’s central place in the understanding of biological diversity. GLOSSARY Evolution. Descent with modification; transformation of species through time, including both changes that occur within species, as well as the origin of new species. Natural Selection. The process in which individuals with a particular trait tend to leave more offspring in the next generation than do individuals with a different trait. Approximately 375 million years ago, a large and vague- ly salamander-like creature plodded from its aquatic home and began the vertebrate invasion of land, setting forththe chain of evolutionary eventsthatled tothe birds thatfillourskies,thebeaststhatwalkoursoil,mewriting this chapter, and you reading it. This was, of course, just one episode in life’s saga: millions of years earlier, plants had come ashore, followed soon thereafter—or perhaps simultaneously—byarthropods.Wecouldgobackmuch earlier, 4 billion years or so, to that fateful day when the first molecule replicated itself, an important milestone in the origin of life and the beginning of the evolutionary pageant. Moving forward, the last few hundred million years have also had their highs and lows: the origins of frogs and trees, the end-Permian extinction when 90 percent of all species perished, and the rise and fall of the dinosaurs. These vignettes are a few of many waypoints in the evolutionary chronicle of life on earth. Evolutionary biologists try to understand this history, explaining how and why life has taken its particular path. But the study of evolution involves more than looking backward to try to understand the past. Evolution is an ongoing process, one possibly operating at a faster rate now than in times past in this human-dominated world. Consequently, evolutionary biology is also forward looking: it includes the study of evolutionary processes in action today— how they operate, what they produce—as well as in- vestigation of how evolution is likely to proceed in the future. Moreover, evolutionary biology is not solely an academic matter; evolution affects humans in many ways, from coping with the emergence of agricultural pests and disease-causing organisms to understanding the workings of our own genome. Indeed, evolutionary
  • 23.
    science hasbroad relevance,playinganimportantrole in advances in many areas, from computer programming to medicine to engineering. 1. WHAT IS EVOLUTION? Lookuptheword“evolution”intheonlineversionofthe Oxford English Dictionary, and you will find 11 defini- tions and numerous subdefinitions, ranging from math- ematical (“the successive transformation of a curve by the alteration of the conditionswhich defineit”) tochem- ical (“the emission or release of gas, heat, light, etc.”) to military (“a manoeuvre executed by troops or ships to adopt a different tactical formation”). Even with ref- erence to biology, there are several definitions, including “emergence or release from an envelope or enclosing structure; (also) protrusion, evagination,” not to men- tion “rare” and “historical” usage related to the concept of preformation of embryos. Even among evolutionary biologists, evolution is defined in different ways. For example, one widely read textbook refers to evolution as “changes in the properties of groups of organisms over the course of generations” (Futuyma 2005), whereas another defines it as “changes in allele frequencies over time” (Freeman and Herron 2007). One might think that—as in so many other areas of evolutionary biology—we could look to Darwin for clarity. But in the first edition of On the Origin of Spe- cies, the term “evolution” never appears (though the last word of the book is “evolved”); not until the sixth edi- tion does Darwin use “evolution.” Rather, Darwin’s term of choice is “descent with modification,” a simple phrase that captures the essence of what evolutionary biology is all about: the study of the transformation of species through time, including both changes that occur within species, as well as the origin of new species. 2. EVOLUTION: PATTERN VERSUS PROCESS Many people—sometimes even biologists—equate evo- lution with natural selection, but the two are not the same. Natural selection is one process that can cause evolutionary change, but natural selection can occur without producing evolutionary change. Conversely, processes other than natural selection can lead to evolution. Natural selection within populations refers to the sit- uation in which individuals with one variant of a trait (say, blue eyes) tend to leave more offspring that are healthy and fertile in the next generation than do in- dividuals with an alternative variant of the trait. Such selection can occur in many ways, for example, if the variantleadstogreaterlongevity,greaterattractivenessto members of the other sex, or greater number of offspring per breeding event. The logic behind natural selection is unassailable. If some trait variant is causally related to greater reproductive success, then more members of the population will have that variant in the next generation; continued over many generations, such selection can greatly change the constitution of a population. But there is a catch. Natural selection can occur with- out leading to evolution if differences among individuals are not genetically based. For natural selection to cause evolutionary change, trait variants must be transmitted from parent to offspring; if that is the case, then offspring will resemble their parents and the trait variants possessed by the parents that produce the most offspring will in- crease in frequency in the next generation. However, offspring do not always resemble their parents. In some cases, individuals vary phenotypically not because they are different genetically, but because they experienced different environments during growth (this is the “nurture” part of the nature versus nurture debate; see chapters III.10 and VII.1). If, in fact, varia- tion in a population is not genetically based, then se- lection will have no evolutionary consequence; in- dividuals surviving and producing many offspring will not differ genetically from those that fail to prosper, and as a result, the gene pool of the population will not change. Nonetheless, much of the phenotypic variation within a population is, in fact, genetically based; con- sequently, natural selection often does lead to evolu- tionary change. But that does not mean that the occurrence of evo- lutionary change necessarily implies the action of nat- ural selection. Other processes—especially mutation, genetic drift, and immigration of individuals with dif- ferent genetic constitutions—also can cause a change in the genetic makeup of a population from one generation to the next (see Section IV: Evolutionary Processes). In other words, natural selection can cause adaptive evo- lutionary change, but not all evolution is adaptive. These caveats notwithstanding, 150 years of research have made clear that naturalselectionisa powerful force responsible for much of the significant evolutionary change that has occurred over the history of life. As the chapters in Section II: Phylogenetics and the History of Life, and Section III: Natural Selection and Adaptation, demonstrate, natural selection can operate in many ways, and scientists have correspondingly devised many methods to detect it, both through studies of the phe- notype and of DNA itself (see also chapter V.14). 3. EVOLUTION: MORE THAN CHANGES IN THE GENE POOL During the heyday of population genetics in the middle decades of the last century, many biologists equated 4 Introduction
  • 24.
    evolution with changesfrom one generation to the next in gene frequencies (gene frequency refers to the fre- quencies of different alleles of a gene; for background on genetic variation, see chapter I.4). The “Modern Synthesis” of the 1930s and 1940s led to several decades in which the field was primarily concerned with the ge- netics of populations with an emphasis on natural se- lection (see chapter I.2). This focus was sharpened by the advent of molecular approaches to studying evolution. Starting in 1960 with the application of enzyme elec- trophoresis techniques, biologists could, for the first time, directly assess the extentof genetic variation within populations. To everyone’s surprise, populations were found to contain much more variation than expected. This finding both challenged the view that natural se- lection was the dominant force guiding evolutionary change (see discussion of “neutralists” in chapters I.2 and V.1), yet further directed attention to the genetics of populations. With more advanced molecular techniques available today, the situation has not changed. There is much more variation than we first suspected. The last 35 years have seen a broadening of evolu- tionary inquiry as the field has recognized that there is more to understanding evolutionary change than study- ing what happens to genes within populations—though this area remains a critically important part of evolu- tionary inquiry. Three aspects of expansion in evolu- tionary thinking are particularly important. First, phenotypic evolution results from evolutionary change in the developmental process that transforms a single-celled fertilized egg into an adult organism. Al- though under genetic control, development is an in- tricate process that cannot be understood by examina- tion of DNA sequences alone. Rather, understanding how phenotypes evolve, and the extent to which devel- opmental systems constrain and direct evolutionary change, requires detailed molecular and embryological knowledge (see chapters V.10 and V.11). Second, history is integral to understanding evolution (see introduction to Section II: Phylogenetics and the History of Life). The study of fossils—paleontology— provides the primary, almost exclusive, direct evidence of life in the past. Somewhat moribund in the middle of the last century, paleontology has experienced a resur- gence in recent decades owing to both dramatic new dis- coveries stemming from an upsurge in paleontological exploration, and new ideas about evolution inspired by and primarily testable with fossil data, such as theories concerning punctuated equilibrium and stasis, species selection, and mass extinction. Initially critical in the development and acceptance of evolutionary theory, paleontology has once again become an important and vibrant part of evolutionary biology (see chapter II.9 and others in Section II). Concurrently, a more fundamental revolution em- phasizing the historical perspective has taken place over the last 30 years with the realization that information on phylogenetic relationships—that is, the tree of life, the pattern of descent and relationship among species—is critical in interpreting all aspects of evolution above the population level. Beginning with a transformation in the field of systematics concerning how phylogenetic re- lationships are inferred, this “tree-thinking” approach now guides study not only of all aspects of macroevolu- tion but also of many population-level phenomena. Finally, life is hierarchically organized. Genes are lo- cated within individuals, individuals within populations, populations within species, and species within clades (a clade consists of an ancestral species and all its de- scendants). Population genetics concerns what happens among individuals within a population, but evolutionary change can occur at all levels. For example, why are there more than 2000 species of rodents but only 3 species of monotremes (the platypus and echidnas), a much older clade of mammals? One cannot look at questions con- cerning natural selection within a population to answer this question. Rather, one must inquire about properties of entire species. Is there some attribute of rodents that makes them particularly prone to speciate or to avoid extinction? Similarly, why is there so much seemingly useless noncoding DNA in the genomes of many species (see chapter V.2)? One possibility is that some genes are particularly adept at mutating to multiply the number of copies of that gene within a genome; such DNA might increase in frequency in the genome even if such multi- plication has no benefit to the individual in whose body the DNA resides. Just as selection among individual or- ganisms on heritable traits can lead to evolutionary change within populations, selection among entities at other levels (species, genes) can also lead to evolutionary change, as long as those entities have traits that are transmitted to their offspring (be they descendant species or genes) and affect the number of descendants they pro- duce.Theupshotisthatevolutionoccursatmultiplelevels of the hierarchy of life; to understand its rich complexity we must study evolution at these distinct levels as well as theinteractionsamongthem.Whathappens,forexample, when a trait that benefits an individual within a popula- tion (perhaps cannibalism—more food, fewer competi- tors!) has detrimental effects at the level of species? Although evolutionary biology has expanded in scope, genetic change is still its fundamental foundation. Nonetheless, in recent years attention has focused on variation that is not genetically based. Phenotypic plas- ticity—the ability of a single genotype to produce dif- ferent phenotypes when exposed to different environ- ments—may itself be adaptive (see chapter III.10). If individuals in a population are likely to experience What Is Evolution? 5
  • 25.
    differentconditionsastheydevelop,thentheevolutionof a genotype thatcould produce appropriate phenotypes depending on circumstances would be advantageous. Although selection on these different phenotypes would not lead to evolutionary change, the degree of plasticity itself can evolve if differences in extent of plasticity lead to differences in the number of surviving offspring. In- deed, an open question is, why don’t populations evolve to become infinitely malleable, capable of producing the appropriatephenotypeforanyenvironment?Presumably, plasticity has an associated cost such that adaptation to different environments often occurs by genetic differ- entiation rather than by the evolution of a single genotype that can produce different phenotypes. Such costs, how- ever, have proven difficult to demonstrate. Differences observed among populations may also reflect plastic responses to different environmental con- ditions and thus may not reflect genetic differentiation. However, if consistently transmitted from one genera- tion to the next, such nongenetic differences may lead to divergent selective pressures on traits that are genetically determined, thus promoting evolutionary divergence between the populations. One particular example con- cerns behavior, which is highly variable in response to the environment—an extreme manifestation of plastic- ity (see chapter VIII.10). Learned behaviors that are transmitted from one generation to the next—often called traditions or culture—occur not only in humans but in other animals, not only our near relatives the apes but also cetaceans, birds, and others. Such behavioral differences among populations would not reflect genetic differentiation, but they might set the stage for genetic divergence in traits relating to the behaviors. One can easily envision, for example, how chimpanzee popula- tions that use different tools—such as delicate twigs to probe termite mounds, or heavy stones to pound nuts— might evolve different morphological features to en- hance the effectiveness of these behaviors. A concrete example involves human populations that tend cattle— surely a nongenetically based behavior—and have evolved genetic changes to permit the digestion of milk in adults. 4. IN THE LIGHT OF EVOLUTION In a 1964 address to the American Society of Zoologists, the distinguished Russian-born biologist Theodosius Dobzhansky proclaimed “nothing makes sense in biol- ogy except in the light of evolution.” Ever since, evolu- tionary biologists have trotted out this phrase (or some permutation of it) to emphasize the centrality of evolu- tion in understanding the biological world. Nonetheless, for much of the twentieth century, the pervasive im- portance of an evolutionary perspective was not at all obvious to many biologists, some of whom considered Dobzhansky’s claim to be self-serving hype. One could argue, for example, that the enormous growth in our understanding of molecular biology from 1950 to 2000 was made with little involvement or insight from evo- lutionary biology. Indeed, to the practicing molecular biologist in the 1980s and 1990s, evolutionary biology was mostly irrelevant. Now, nothing could be further from the truth. When results of the human genome sequencing project first appeared in2000, many initiallybelieved that a thorough understanding of human biology would soon follow, answering questions about the genetic basis of human diseases and phenotypic variation among individuals. These hopes were quickly dashed—the genetic code,after all, is nothing more than a long list of letters (A, C, G, and T, the abbreviations of the four nucleotide building blocks of DNA). Much of the genome of many species seems to have no function and is just, in some sense, functionless filler; as a result, picking out where the genes lieinthis4billion–longstringofalphabetspaghetti,much less figuring out how these genes function, is not easy. Sowhere did molecular biologists turn? Tothe fieldof evolutionary biology! Genomicists soon realized that the best way to understand the human genome was to study it in the context of its evolutionary history, by comparing human sequences with those of other species in a phy- logenetic framework. One method for locating genes, for example, is to examine comparable parts of the genome of different species. The underlying rationale is that genes evolve more slowly than other parts of the genome. Specifically, nonfunctioning stretches of DNA tend to evolve differences through time as random mu- tations become established (the process of genetic drift; see chapter IV.1), but functioning genes tend to diverge less, because natural selection removes deleterious mu- tations when they arise, keeping the DNA sequence sim- ilar among species. As a result, examination of the amount of divergence between two species relative to the amountof time since theyshareda commonancestor can pinpoint stretches of DNA where evolution has occurred slowly, thus identifying the position of functional genes. Moreover, how a gene functions can often be deduced by comparing its function with that of homologous genes in other species and using a phylogeny to reconstruct the gene’s evolutionary history (see chapter V.14). And thus was born the effort to sequence the ge- nomes of other species (see chapter V.3). At first, the nascent field of comparative genomics focused on pri- mates and model laboratory species such as mice and fruit flies, the former to permit comparisons of the human genome with that of our close evolutionary rel- atives, the latter to take advantage of the great under- standing of the genomic systems of well-studied species. 6 Introduction
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    CHAPTER XXIV. THE REASONAND END OF DIVINE PUNISHMENTS. Now that is in our power, of which equally with its opposite we are masters,—as, say, to philosophize or not, to believe or disbelieve. In consequence, then, of our being equally masters of each of the opposites, what depends on us is found possible. Now the commandments may be done or not done by us, who, as is reasonable, are liable to praise and blame. And those, again, who are punished on account of sins committed by them, are punished for them alone; for what is done is past, and what is done can never be undone. The sins committed before faith are accordingly forgiven by the Lord, not that they may be undone, but as if they had not been done. “But not all,” says Basilides, “but only sins involuntary and in ignorance, are forgiven;” as would be the case were it a man, and not God, that conferred such a boon. To such an one Scripture says, “Thou thoughtest that I would be like thee.”[662] But if we are punished for voluntary sins, we are punished not that the sins which are done may be undone, but because they were done. But punishment does not avail to him who has sinned, to undo his sin, but that he may sin no more, and that no one else fall into the like. Therefore the good God corrects for these three causes: First, that he who is corrected may become better than his former self; then that those who are capable of being saved by examples may be driven back, being admonished; and thirdly, that he who is injured may not be readily despised, and be apt to receive injury. And there are two methods of correction—the instructive and the punitive, which we have called the disciplinary. It ought to be known, then, that those who fall into sin after baptism[663] are those who are subjected to discipline; for the deeds done before are remitted, and those done after are purged. It is in reference to the unbelieving that it is said, “that they are reckoned as the chaff which the wind drives from the face of the earth, and the drop which falls from a vessel.”[664]
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    CHAPTER XXV. TRUE PERFECTIONCONSISTS IN THE KNOWLEDGE AND LOVE OF GOD. “Happy he who possesses the culture of knowledge, and is not moved to the injury of the citizens or to wrong actions, but contemplates the undecaying order of immortal nature, how and in what way and manner it subsists. To such the practice of base deeds attaches not.” Rightly, then, Plato says, “that the man who devotes himself to the contemplation of ideas will live as a god among men; now the mind is the place of ideas, and God is mind.” He says that he who contemplates the unseen God lives as a god among men. And in the Sophist, Socrates calls the stranger of Elea, who was a dialectician, “god:” “Such are the gods who, like stranger guests, frequent cities. For when the soul, rising above the sphere of generation, is by itself apart, and dwells amidst ideas,” like the Coryphæus in Theætetus, now become as an angel, it will be with Christ, being rapt in contemplation, ever keeping in view the will of God; in reality “Alone wise, while these flit like shadows.”[665] “For the dead bury their dead.” Whence Jeremiah says: “I will fill it with the earth-born dead whom mine anger has smitten.”[666] God, then, being not a subject for demonstration, cannot be the object of science. But the Son is wisdom, and knowledge, and truth, and all else that has affinity thereto. He is also susceptible of demonstration and of description. And all the powers of the Spirit, becoming collectively one thing, terminate in the same point—that is, in the Son. But He is incapable of being declared, in respect of the idea of each one of His powers. And the Son is neither simply one thing as one thing, nor many things as parts, but one thing as all things; whence also He is all things. For He is the circle of all powers rolled and united into one unity. Wherefore the Word is called the Alpha and the Omega, of whom alone the end becomes
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    beginning, and endsagain at the original beginning without any break. Wherefore also to believe in Him, and by Him, is to become a unit, being indissolubly united in Him; and to disbelieve is to be separated, disjoined, divided. “Wherefore thus saith the Lord, Every alien son is uncircumcised in heart, and uncircumcised in flesh” (that is, unclean in body and soul): “there shall not enter one of the strangers into the midst of the house of Israel, but the Levites.”[667] He calls those that would not believe, but would disbelieve, strangers. Only those who live purely being true priests of God. Wherefore, of all the circumcised tribes, those anointed to be high priests, and kings, and prophets, were reckoned more holy. Whence He commands them not to touch dead bodies, or approach the dead; not that the body was polluted, but that sin and disobedience were incarnate, and embodied, and dead, and therefore abominable. It was only, then, when a father and mother, a son and daughter died, that the priest was allowed to enter, because these were related only by flesh and seed, to whom the priest was indebted for the immediate cause of his entrance into life. And they purify themselves seven days, the period in which Creation was consummated. For on the seventh day the rest is celebrated; and on the eighth he brings a propitiation, as is written in Ezekiel, according to which propitiation the promise is to be received.[668] And the perfect propitiation, I take it, is that propitious faith in the gospel which is by the law and the prophets, and the purity which shows itself in universal obedience, with the abandonment of the things of the world; in order to that grateful surrender of the tabernacle, which results from the enjoyment of the soul. Whether, then, the time be that which through the seven periods enumerated returns to the chiefest rest,[669] or the seven heavens, which some reckon one above the other; or whether also the fixed sphere which borders on the intellectual world be called the eighth, the expression denotes that the Gnostic ought to rise out of the sphere of creation and of sin. After these seven days, sacrifices are offered for sins. For there is still fear of change, and it touches
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    the seventh circle.The righteous Job says: “Naked came I out of my mother’s womb, and naked shall I return there;”[670] not naked of possessions, for that were a trivial and common thing; but, as a just man, he departs naked of evil and sin, and of the unsightly shape which follows those who have led bad lives. For this was what was said, “Unless ye be converted, and become as children,”[671] pure in flesh, holy in soul by abstinence from evil deeds; showing that He would have us to be such as also He generated us from our mother —the water.[672] For the intent of one generation succeeding another is to immortalize by progress. “But the lamp of the wicked shall be put out.”[673] That purity in body and soul which the Gnostic partakes of, the all-wise Moses indicated, by employing repetition in describing the incorruptibility of body and of soul in the person of Rebecca, thus: “Now the virgin was fair, and man had not known her.”[674] And Rebecca, interpreted, means “glory of God;” and the glory of God is immortality. This is in reality righteousness, not to desire other things, but to be entirely the consecrated temple of the Lord. Righteousness is peace of life and a well-conditioned state, to which the Lord dismissed her when He said, “Depart into peace.”[675] For Salem is, by interpretation, peace; of which our Saviour is enrolled King, as Moses says, Melchizedek king of Salem, priest of the most high God, who gave bread and wine, furnishing consecrated food for a type of the Eucharist. And Melchizedek is interpreted “righteous king;” and the name is a synonym for righteousness and peace. Basilides, however, supposes that Righteousness and her daughter Peace dwell stationed in the eighth sphere. But we must pass from physics to ethics, which are clearer; for the discourse concerning these will follow after the treatise in hand. The Saviour Himself, then, plainly initiates us into the mysteries, according to the words of the tragedy:[676] “Seeing those who see, he also gives the orgies.”
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    And if youask, “These orgies, what is their nature?” You will hear again: “It is forbidden to mortals uninitiated in the Bacchic rites to know.” And if any one will inquire curiously what they are, let him hear: “It is not lawful for thee to hear, but they are worth knowing; The rites of the God detest him who practises impiety.” Now God, who is without beginning, is the perfect beginning of the universe, and the producer of the beginning. As, then, He is being, He is the first principle of the department of action, as He is good, of morals; as He is mind, on the other hand, He is the first principle of reasoning and of judgment. Whence also He alone is Teacher, who is the only Son of the Most High Father, the Instructor of men.
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    CHAPTER XXVI. HOW THEPERFECT MAN TREATS THE BODY AND THE THINGS OF THE WORLD. Those, then, who run down created existence and vilify the body are wrong; not considering that the frame of man was formed erect for the contemplation of heaven, and that the organization of the senses tends to knowledge; and that the members and parts are arranged for good, not for pleasure. Whence this abode becomes receptive of the soul which is most precious to God; and is dignified with the Holy Spirit through the sanctification of soul and body, perfected with the perfection of the Saviour. And the succession of the three virtues is found in the Gnostic, who morally, physically, and logically occupies himself with God. For wisdom is the knowledge of things divine and human; and righteousness is the concord of the parts of the soul; and holiness is the service of God. But if one were to say that he disparaged the flesh, and generation on account of it, by quoting Isaiah, who says, “All flesh is grass, and all the glory of man as the flower of grass: the grass is withered, and the flower has fallen; but the word of the Lord endureth for ever;”[677] let him hear the Spirit interpreting the matter in question by Jeremiah, “And I scattered them like dry sticks, that are made to fly by the wind into the desert. This is the lot and portion of your disobedience, saith the Lord. As thou hast forgotten me, and hast trusted in lies, so will I discover thy hinder parts to thy face; and thy disgrace shall be seen, thy adultery, and thy neighing,” and so on.[678] For “the flower of grass,” and “walking after the flesh,” and “being carnal,” according to the apostle, are those who are in their sins. The soul of man is confessedly the better part of man, and the body the inferior. But neither is the soul good by nature, nor, on the other hand, is the body bad by nature. Nor is that which is not good straightway bad. For there are things which occupy a middle place, and among them are things to be preferred, and things to be rejected. The constitution of man, then, which has its place among things of
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    sense, was necessarilycomposed of things diverse, but not opposite —body and soul. Always therefore the good actions, as better, attach to the better and ruling spirit; and voluptuous and sinful actions are attributed to the worse, the sinful one. Now the soul of the wise man and Gnostic, as sojourning in the body, conducts itself towards it gravely and respectfully, not with inordinate affections, as about to leave the tabernacle if the time of departure summon. “I am a stranger in the earth, and a sojourner with you,” it is said.[679] And hence Basilides says, that he apprehends that the election are strangers to the world, being supramundane by nature. But this is not the case. For all things are of one God. And no one is a stranger to the world by nature, their essence being one, and God one. But the elect man dwells as a sojourner, knowing all things to be possessed and disposed of; and he makes use of the things which the Pythagoreans make out to be the threefold good things. The body, too, as one sent on a distant pilgrimage, uses inns and dwellings by the way, having care of the things of the world, of the places where he halts; but leaving his dwelling-place and property without excessive emotion; readily following him that leads him away from life; by no means and on no occasion turning back; giving thanks for his sojourn, and blessing [God] for his departure, embracing the mansion that is in heaven. “For we know, that, if the earthly house of our tabernacle be dissolved, we have a building of God, an house not made with hands, eternal in the heavens. For we that are in this tabernacle do groan, desiring to be clothed upon with our house which is from heaven: if so be that being clothed we shall not be found naked. For we walk by faith, not by sight,”[680] as the apostle says; “and we are willing rather to be absent from the body, and present with God.” The rather is in comparison. And comparison obtains in the case of things that fall under resemblance; as the more valiant man is more valiant among the valiant, and most valiant among cowards. Whence he adds, “Wherefore we strive, whether present or absent, to be
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    accepted with Him,”[681]that is, God, whose work and creation are all things, both the world and things supramundane. I admire Epicharmus, who clearly says: “Endowed with pious mind, you will not, in dying, Suffer aught evil. The spirit will dwell in heaven above;” and the minstrel[682] who sings: “The souls of the wicked flit about below the skies on earth, In murderous pains beneath inevitable yokes of evils; But those of the pious dwell in the heavens, Hymning in songs the Great, the Blessed One.” The soul is not then sent down from heaven to what is worse. For God works all things up to what is better. But the soul which has chosen the best life—the life that is from God and righteousness— exchanges earth for heaven. With reason therefore, Job, who had attained to knowledge, said, “Now I know that Thou canst do all things; and nothing is impossible to Thee. For who tells me of what I know not, great and wonderful things with which I was unacquainted? And I felt myself vile, considering myself to be earth and ashes.”[683] For he who, being in a state of ignorance, is sinful, “is earth and ashes;” while he who is in a state of knowledge, being assimilated as far as possible to God, is already spiritual, and so elect. And that Scripture calls the senseless and disobedient “earth,” will be made clear by Jeremiah the prophet, saying, in reference to Joachim and his brethren, “Earth, earth, hear the word of the Lord; Write this man, as man excommunicated.”[684] And another prophet says again, “Hear, O heaven; and give ear, O earth,”[685] calling understanding “ear,” and the soul of the Gnostic, that of the man who has applied himself to the contemplation of heaven and divine things, and in this way has become an Israelite, “heaven.” For again he calls him who has made ignorance and hardness of heart his choice, “earth.” And the expression “give ear” he derives from the “organs of hearing,” “the ears,” attributing carnal things to those
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    who cleave tothe things of sense. Such are they of whom Micah the prophet says, “Hear the word of the Lord, ye peoples who dwell with pangs.”[686] And Abraham said, “By no means. The Lord is He who judgeth the earth;”[687] “since he that believeth not, is,” according to the utterance of the Saviour, “condemned already.”[688] And there is written in the Kings[689] the judgment and sentence of the Lord, which stands thus: “The Lord hears the righteous, but the wicked He saveth not, because they do not desire to know God.” For the Almighty will not accomplish what is absurd. What do the heresies say to this utterance, seeing Scripture proclaims the Almighty God to be good, and not the author of evil and wrong, if indeed ignorance arises from one not knowing? But God does nothing absurd. “For this God,” it is said, “is our God, and there is none to save besides Him.”[690] “For there is no unrighteousness with God,”[691] according to the apostle. And clearly yet the prophet teaches the will of God, and the gnostic proficiency, in these words: “And now, Israel, what doth the Lord God require of thee, but to fear the Lord thy God, and walk in all His ways, and love Him, and serve Him alone?”[692] He asks of thee, who hast the power of choosing salvation. What is it, then, that the Pythagoreans mean when they bid us “pray with the voice?” As seems to me, not that they thought the Divinity could not hear those who speak silently, but because they wished prayers to be right, which no one would be ashamed to make in the knowledge of many. We shall, however, treat of prayer in due course by and by. But we ought to have works that cry aloud, as becoming “those who walk in the day.”[693] “Let thy works shine,”[694] and behold a man and his works before his face. “For behold God and His works.”[695] For the gnostic must, as far as is possible, imitate God. And the poets call the elect in their pages godlike and gods, and equal to the gods, and equal in sagacity to Zeus, and having counsels like the gods, and resembling the gods,—nibbling, as seems to me, at the expression, “in the image and likeness.”[696]
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    Euripides accordingly says,“Golden wings are round my back, and I am shod with the winged sandals of the Sirens; and I shall go aloft into the wide ether, to hold converse with Zeus.” But I shall pray the Spirit of Christ to wing me to my Jerusalem. For the Stoics say that heaven is properly a city, but places here on earth are not cities; for they are called so, but are not. For a city is an important thing, and the people a decorous body, and a multitude of men regulated by law as the church by the word—a city on earth impregnable—free from tyranny; a product of the divine will on earth as in heaven. Images of this city the poets create with their pen. For the Hyperboreans, and the Arimaspian cities, and the Elysian plains, are commonwealths of just men. And we know Plato’s city placed as a pattern in heaven.
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    BOOK V. CHAPTER I. ONFAITH. f the gnostic so much has been cursorily, as it were, written. We proceed now to the sequel, and must again contemplate faith; for there are some that draw the distinction, that faith has reference to the Son, and knowledge to the Spirit. But it has escaped their notice that, in order to believe truly in the Son, we must believe that He is the Son, and that He came, and how, and for what, and respecting His passion; and we must know who is the Son of God. Now neither is knowledge without faith, nor faith without knowledge. Nor is the Father without the Son; for the Son is with the Father. And the Son is the true teacher respecting the Father; and that we may believe in the Son, we must know the Father, with whom also is the Son. Again, in order that we may know the Father, we must believe in the Son, that it is the Son of God who teaches; for from faith to knowledge by the Son is the Father. And the knowledge of the Son and Father, which is according to the gnostic rule—that which in reality is gnostic—is the attainment and comprehension of the truth by the truth. We, then, are those who are believers in what is not believed, and who are gnostics as to what is unknown; that is, gnostics as to what is unknown and disbelieved by all, but believed and known by a few; and gnostics, not describing actions by speech, but gnostics in the exercise of contemplation. Happy is he who speaks in the ears of the hearing. Now Faith is the ear of the soul. And such the Lord intimates faith to be, when He says, “He that hath ears to hear, let him hear;”[697] so that by believing he may comprehend what He
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    says, as Hesays it. Homer, too, the oldest of the poets, using the word “hear” instead of “perceive”—the specific for the generic term —writes: “Him most they heard.”[698] For, in fine, the agreement and harmony of the faith of both[699] contribute to one end—salvation. We have in the apostle an unerring witness: “For I desire to see you, that I may impart unto you some spiritual gift, in order that ye may be strengthened; that is, that I may be comforted in you, by the mutual faith of you and me.”[700] And further on again he adds, “The righteousness of God is revealed from faith to faith.”[701] The apostle, then, manifestly announces a double faith, or rather one which admits of growth and perfection; for the common faith lies beneath as a foundation. To those, therefore, who desire to be healed, and are moved by faith, He added, “Thy faith hath saved thee.”[702] But that which is excellently built upon is consummated in the believer, and is again perfected by the faith which results from instruction and the word, in order to the performance of the commandments. Such were the apostles, in whose case it is said that “faith removed mountains and transplanted trees.”[703] Whence, perceiving the greatness of its power, they asked “that faith might be added to them;”[704] a faith which salutarily bites the soil “like a grain of mustard,” and grows magnificently in it, to such a degree that the reasons of things sublime rest on it. For if one by nature knows God, as Basilides thinks, who calls intelligence of a superior order at once faith and kingship, and a creation worthy of the essence of the Creator; and explains that near Him exists not power, but essence and nature and substance; and says that faith is not the rational assent of the soul exercising free-will, but an undefined beauty, belonging immediately to the creature;—the precepts both of the Old and of the New Testament are, then, superfluous, if one is saved by nature, as Valentinus would have it, and is a believer and an elect man by nature, as Basilides thinks; and nature would have been able, one
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    time or other,to have shone forth, apart from the Saviour’s appearance. But were they to say that the visit of the Saviour was necessary, then the properties of nature are gone from them, the elect being saved by instruction, and purification, and the doing of good works. Abraham, accordingly, who through hearing believed the voice, which promised under the oak in Mamre, “I will give this land to thee, and to thy seed,” was either elect or not. But if he was not, how did he straightway believe, as it were naturally? And if he was elect, their hypothesis is done away with, inasmuch as even previous to the coming of the Lord an election was found, and that saved: “For it was reckoned to him for righteousness.”[705] For if any one, following Marcion, should dare to say that the Creator (Δημιουργόν) saved the man that believed on him, even before the advent of the Lord, (the election being saved with their own proper salvation); the power of the good Being will be eclipsed; inasmuch as late only, and subsequent to the Creator spoken of by them in words of good omen, it made the attempt to save, and by his instruction, and in imitation of him. But if, being such, the good Being save, according to them; neither is it his own that he saves, nor is it with the consent of him who formed the creation that he essays salvation, but by force or fraud. And how can he any more be good, acting thus, and being posterior? But if the locality is different, and the dwelling-place of the Omnipotent is remote from the dwelling-place of the good God; yet the will of him who saves, having been the first to begin, is not inferior to that of the good God. From what has been previously proved, those who believe not are proved senseless: “For their paths are perverted, and they know not peace,” saith the prophet.[706] “But foolish and unlearned questions” the divine Paul exhorted to “avoid, because they gender strifes.”[707] And Æschylus exclaims: “In what profits not, labour not in vain.” For that investigation, which accords with faith, which builds, on the foundation of faith, the august knowledge of the truth, we know to be the best. Now we know that neither things which are clear are
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    made subjects ofinvestigation, such as if it is day, while it is day; nor things unknown, and never destined to become clear, as whether the stars are even or odd in number; nor things convertible; and those are so which can be said equally by those who take the opposite side, as if what is in the womb is a living creature or not. A fourth mode is, when, from either side of those, there is advanced an unanswerable and irrefragable argument. If, then, the ground of inquiry, according to all of these modes, is removed, faith is established. For we advance to them the unanswerable consideration, that it is God who speaks and comes to our help in writing, respecting each one of the points regarding which I investigate. Who, then, is so impious as to disbelieve God, and to demand proofs from God as from men? Again, some questions demand the evidence of the senses, as if one were to ask whether the fire be warm, or the snow white; and some admonition and rebuke, as the question if you ought to honour your parents. And there are those that deserve punishment, as to ask proofs of the existence of Providence. There being then a Providence, it were impious to think that the whole of prophecy and the economy in reference to a Saviour did not take place in accordance with Providence. And perchance one should not even attempt to demonstrate such points, the divine Providence being evident from the sight of all its skilful and wise works which are seen, some of which take place in order, and some appear in order. And He who communicated to us being and life, has communicated to us also reason, wishing us to live rationally and rightly. For the Word of the Father of the universe is not the uttered word (λόγος προφορικός), but the wisdom and most manifest kindness of God, and His power too, which is almighty and truly divine, and not incapable of being conceived by those who do not confess—the all-potent will. But since some are unbelieving, and some are disputatious, all do not attain to the perfection of the good. For neither is it possible to attain it without the exercise of free choice; nor does the whole depend on our own purpose; as, for example, what is destined to happen. “For by grace we are saved:” not, indeed, without good works; but we must, by being formed for what is good, acquire an
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    inclination for it.And we must possess the healthy mind which is fixed on the pursuit of the good; in order to which we have the greatest need of divine grace, and of right teaching, and of holy susceptibility, and of the drawing of the Father to Him. For, bound in this earthly body, we apprehend the objects of sense by means of the body; but we grasp intellectual objects by means of the logical faculty itself. But if one expect to apprehend all things by the senses, he has fallen far from the truth. Spiritually, therefore, the apostle writes respecting the knowledge of God, “For now we see as through a glass, but then face to face.”[708] For the vision of the truth is given but to few. Accordingly, Plato says in the Epinomis, “I do not say that it is possible for all to be blessed and happy; only a few. Whilst we live, I pronounce this to be the case. But there is a good hope that after death I shall attain all.” To the same effect is what we find in Moses: “No man shall see my face, and live.”[709] For it is evident that no one during the period of life has been able to apprehend God clearly. But “the pure in heart shall see God,”[710] when they arrive at the final perfection. For since the soul became too enfeebled for the apprehension of realities, we needed a divine teacher. The Saviour is sent down—a teacher and leader in the acquisition of the good—the secret and sacred token of the great Providence. “Where, then, is the scribe? where is the searcher of this world? Hath not God made foolish the wisdom of this world?”[711] it is said. And again, “I will destroy the wisdom of the wise, and bring to nothing the understanding of the prudent,”[712] plainly of those wise in their own eyes, and disputatious. Excellently therefore Jeremiah says, “Thus saith the Lord, Stand in the ways, and ask for the eternal paths, what is the good way, and walk in it, and ye shall find expiation for your souls.”[713] Ask, he says, and inquire of those who know, without contention and dispute. And on learning the way of truth, let us walk on the right way, without turning till we attain to what we desire. It was therefore with reason that the king of the Romans (his name was Numa), being a Pythagorean, first of all men, erected a temple to Faith and Peace. “And to Abraham, on believing,
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    righteousness was reckoned.”[714]He, prosecuting the lofty philosophy of aerial phenomena, and the sublime philosophy of the movements in the heavens, was called Abram, which is interpreted “sublime father.”[715] But afterwards, on looking up to heaven, whether it was that he saw the Son in the spirit, as some explain, or a glorious angel, or in any other way recognised God to be superior to the creation, and all the order in it, he receives in addition the Alpha, the knowledge of the one and only God, and is called Abraam, having, instead of a natural philosopher, become wise, and a lover of God. For it is interpreted, “elect father of sound.” For by sound is the uttered word: the mind is its father; and the mind of the good man is elect. I cannot forbear praising exceedingly the poet of Agrigentum, who celebrates faith as follows: “Friends, I know, then, that there is truth in the myths Which I will relate. But very difficult to men, And irksome to the mind, is the attempt of faith.”[716] Wherefore also the apostle exhorts, “that your faith should not be in the wisdom of men,” who profess to persuade, “but in the power of God,”[717] which alone without proofs, by mere faith, is able to save. “For the most approved of those that are reputable knows how to keep watch. And justice will apprehend the forgers and witnesses of lies,” says the Ephesian.[718] For he, having derived his knowledge from the barbarian philosophy, is acquainted with the purification by fire of those who have led bad lives, which the Stoics afterwards called the Conflagration (ἐκπύρωσις), in which also they teach that each will arise exactly as he was, so treating of the resurrection; while Plato says as follows, that the earth at certain periods is purified by fire and water: “There have been many destructions of men in many ways; and there shall be very great ones by fire and water; and others briefer by innumerable causes.” And after a little he adds: “And, in truth, there is a change of the objects which revolve about earth and heaven; and in the course of long periods there is the destruction of the objects on earth by a great conflagration.” Then he subjoins respecting the deluge: “But when,
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    again, the godsdeluge the earth to purify it with water, those on the mountains, herdsmen and shepherds, are saved; those in your cities are carried down by the rivers into the sea.” And we showed in the first Miscellany that the philosophers of the Greeks are called thieves, inasmuch as they have taken without acknowledgment their principal dogmas from Moses and the prophets. To which also we shall add, that the angels who had obtained the superior rank, having sunk into pleasures, told to the women the secrets which had come to their knowledge; while the rest of the angels concealed them, or rather, kept them against the coming of the Lord. Thence emanated the doctrine of providence, and the revelation of high things; and prophecy having already been imparted to the philosophers of the Greeks, the treatment of dogma arose among the philosophers, sometimes true when they hit the mark, and sometimes erroneous, when they comprehended not the secret of the prophetic allegory. And this it is proposed briefly to indicate in running over the points requiring mention. Faith, then, we say, we are to show must not be inert and alone, but accompanied with investigation. For I do not say that we are not to inquire at all. For “Search, and thou shalt find,”[719] it is said. “What is sought may be captured, But what is neglected escapes,” according to Sophocles. The like also says Menander the comic poet: “All things sought, The wisest say, need anxious thought.” But we ought to direct the visual faculty of the soul aright to discovery, and to clear away obstacles; and to cast clean away contention, and envy, and strife, destined to perish miserably from among men. For very beautifully does Timon of Phlius write:
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    “And Strife, thePlague of Mortals, stalks vainly shrieking, The sister of Murderous Quarrel and Discord, Which rolls blindly over all things. But then It sets its head towards men, and casts them on hope.” Then a little below he adds: “For who hath set these to fight in deadly strife? A rabble keeping pace with Echo; for, enraged at those silent, It raised an evil disease against men, and many perished;” of the speech which denies what is false, and of the dilemma, of that which is concealed, of the Sorites, and of the Crocodilean, of that which is open, and of ambiguities and sophisms. To inquire, then, respecting God, if it tend not to strife, but to discovery, is salutary. For it is written in David, “The poor eat, and shall be filled; and they shall praise the Lord that seek Him. Your heart shall live for ever.”[720] For they who seek Him after the true search, praising the Lord, shall be filled with the gift that comes from God, that is, knowledge. And their soul shall live; for the soul is figuratively termed the heart, which ministers life: for by the Son is the Father known. We ought not to surrender our ears to all who speak and write rashly. For cups also, which are taken hold of by many by the ears, are dirtied, and lose the ears; and besides, when they fall they are broken. In the same way also, those, who have polluted the pure hearing of faith by many trifles, at last becoming deaf to the truth, become useless and fall to the earth. It is not, then, without reason that we commanded boys to kiss their relations, holding them by the ears; indicating this, that the feeling of love is engendered by hearing. And “God,” who is known to those who love, “is love,”[721] as “God,” who by instruction is communicated to the faithful, “is faithful;”[722] and we must be allied to Him by divine love: so that by like we may see like, hearing the word of truth guilelessly and purely, as children who obey us. And this was what he, whoever he
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    was, indicated whowrote on the entrance to the temple at Epidaurus the inscription: “Pure he must be who goes within The incense-perfumed fane.” And purity is “to think holy thoughts.” “Except ye become as these little children, ye shall not enter,” it is said, “into the kingdom of heaven.”[723] For there the temple of God is seen established on three foundations—faith, hope, and love.
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    CHAPTER II. ON HOPE. Respectingfaith we have adduced sufficient testimonies of writings among the Greeks. But in order not to exceed bounds, through eagerness to collect a very great many also respecting hope and love, suffice it merely to say that in the Crito Socrates, who prefers a good life and death to life itself, thinks that we have hope of another life after death. Also in the Phædrus he says, “That only when in a separate state can the soul become partaker of the wisdom which is true, and surpasses human power; and when, having reached the end of hope by philosophic love, desire shall waft it to heaven, then,” says he, “does it receive the commencement of another, an immortal life.” And in the Symposium he says, “That there is instilled into all the natural love of generating what is like, and in men of generating men alone, and in the good man of the generation of the counterpart of himself. But it is impossible for the good man to do this without possessing the perfect virtues, in which he will train the youth who have recourse to him.” And as he says in the Theætetus, “He will beget and finish men. For some procreate by the body, others by the soul;” since also with the barbarian philosophers to teach and enlighten is called to regenerate; and “I have begotten you in Jesus Christ,”[724] says the good apostle somewhere. Empedocles, too, enumerates friendship among the elements, conceiving it as a combining love: “Which do you look at with your mind; and don’t sit gaping with your eyes.” Parmenides, too, in his poem, alluding to hope, speaks thus: “Yet look with the mind certainly on what is absent as present, For it will not sever that which is from the grasp it has of that which is Not, even if scattered in every direction over the world or combined.”
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    CHAPTER III. THE OBJECTSOF FAITH AND HOPE PERCEIVED BY THE MIND ALONE. For he who hopes, as he who believes, sees intellectual objects and future things with the mind. If, then, we affirm that aught is just, and affirm it to be good, and we also say that truth is something, yet we have never seen any of such objects with our eyes, but with our mind alone. Now the Word of God says, “I am the truth.”[725] The Word is then to be contemplated by the mind. “Do you aver,” it was said,[726] “that there are any true philosophers?” “Yes,” said I, “those who love to contemplate the truth.” In the Phædrus also, Plato, speaking of the truth, shows it as an idea. Now an idea is a conception of God; and this the barbarians have termed the Word of God. The words are as follow: “For one must then dare to speak the truth, especially in speaking of the truth. For the essence of the soul, being colourless, formless, and intangible, is visible only to God,[727] its guide.” Now the Word issuing forth was the cause of creation; then also he generated himself, “when the Word had become flesh,”[728] that He might be seen. The righteous man will seek the discovery that flows from love, to which if he hastes he prospers. For it is said, “To him that knocketh, it shall be opened: ask, and it shall be given to you.”[729] “For the violent that storm the kingdom”[730] are not so in disputatious speeches; but by continuance in a right life and unceasing prayers, are said “to take it by force,” wiping away the blots left by their previous sins. “You may obtain wickedness, even in great abundance.[731] And him who toils God helps; For the gifts of the Muses, hard to win, Lie not before you, for any one to bear away.” The knowledge of ignorance is, then, the first lesson in walking according to the Word. An ignorant man has sought, and having sought, he finds the teacher; and finding has believed, and believing has hoped; and henceforward having loved, is assimilated to what
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    was loved—endeavouring tobe what he first loved. Such is the method Socrates shows Alcibiades, who thus questions: “Do you not think that I shall know about what is right otherwise?” “Yes, if you have found out.” “But you don’t think I have found out?” “Certainly, if you have sought.” “Then you don’t think that I have sought?” “Yes, if you think you do not know.”[732] So with the lamps of the wise virgins, lighted at night in the great darkness of ignorance, which the Scripture signified by “night.” Wise souls, pure as virgins, understanding themselves to be situated amidst the ignorance of the world, kindle the light, and rouse the mind, and illumine the darkness, and dispel ignorance, and seek truth, and await the appearance of the Teacher. “The mob, then,” said I, “cannot become a philosopher.”[733] “Many rod-bearers there are, but few Bacchi,” according to Plato. “For many are called, but few chosen.”[734] “Knowledge is not in all,”[735] says the apostle. “And pray that we may be delivered from unreasonable and wicked men: for all men have not faith.”[736] And the Poetics of Cleanthes, the Stoic, writes to the following effect: “Look not to glory, wishing to be suddenly wise, And fear not the undiscerning and rash opinion of the many; For the multitude has not an intelligent, or wise, or right judgment, And it is in few men that you will find this.”[737] And more sententiously the comic poet briefly says: “It is a shame to judge of what is right by much noise.” For they heard, I think, that excellent wisdom, which says to us, “Watch your opportunity in the midst of the foolish, and in the midst of the intelligent continue.”[738] And again, “The wise will conceal sense.”[739] For the many demand demonstration as a pledge of truth, not satisfied with the bare salvation by faith.
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    “But it isstrongly incumbent to disbelieve the dominant wicked, And as is enjoined by the assurance of our muse, Know by dissecting the utterance within your breast.” “For this is habitual to the wicked,” says Empedocles, “to wish to overbear what is true by disbelieving it.” And that our tenets are probable and worthy of belief, the Greeks shall know, the point being more thoroughly investigated in what follows. For we are taught what is like by what is like. For says Solomon, “Answer a fool according to his folly.”[740] Wherefore also, to those that ask the wisdom that is with us, we are to hold out things suitable, that with the greatest possible ease they may, through their own ideas, be likely to arrive at faith in the truth. For “I became all things to all men, that I might gain all men.”[741] Since also “the rain” of the divine grace is sent down “on the just and the unjust.”[742] “Is He the God of the Jews only, and not also of the Gentiles? Yes, also of the Gentiles: if indeed He is one God,”[743] exclaims the noble apostle.
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    CHAPTER IV. DIVINE THINGSWRAPPED UP IN FIGURES BOTH IN THE SACRED AND IN HEATHEN WRITERS. But since they will believe neither in what is good justly nor in knowledge unto salvation, we ourselves reckoning what they claim as belonging to us, because all things are God’s; and especially since what is good proceeded from us to the Greeks, let us handle those things as they are capable of hearing. For intelligence or rectitude this great crowd estimates not by truth, but by what they are delighted with. And they will be pleased not more with other things than with what is like themselves. For he who is still blind and dumb, not having understanding, or the undazzled and keen vision of the contemplative soul, which the Saviour confers, like the uninitiated at the mysteries, or the unmusical at dances, not being yet pure and worthy of the pure truth, but still discordant and disordered and material, must stand outside of the divine choir. “For we compare spiritual things with spiritual.”[744] Wherefore, in accordance with the method of concealment, the truly sacred Word, truly divine and most necessary for us, deposited in the shrine of truth, was by the Egyptians indicated by what were called among them adyta, and by the Hebrews by the veil. Only the consecrated—that is, those devoted to God, circumcised in the desires of the passions for the sake of love to that which is alone divine—were allowed access to them. For Plato also thought it not lawful for “the impure to touch the pure.” Thence the prophecies and oracles are spoken in enigmas, and the mysteries are not exhibited incontinently to all and sundry, but only after certain purifications and previous instructions. “For the Muse was not then Greedy of gain or mercenary; Nor were Terpsichore’s sweet, Honey-toned, silvery soft-voiced Strains made merchandise of.”
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    Now those instructedamong the Egyptians learned first of all that style of the Egyptian letters which is called Epistolographic; and second, the Hieratic, which the sacred scribes practise; and finally, and last of all, the Hieroglyphic, of which one kind which is by the first elements is literal (Kyriologic), and the other Symbolic. Of the Symbolic, one kind speaks literally by imitation, and another writes as it were figuratively; and another is quite allegorical, using certain enigmas. Wishing to express Sun in writing, they make a circle; and Moon, a figure like the Moon, like its proper shape. But in using the figurative style, by transposing and transferring, by changing and by transforming in many ways as suits them, they draw characters. In relating the praises of the kings in theological myths, they write in anaglyphs.[745] Let the following stand as a specimen of the third species—the Enigmatic. For the rest of the stars, on account of their oblique course, they have figured like the bodies of serpents; but the sun like that of a beetle, because it makes a round figure of ox- dung, and rolls it before its face. And they say that this creature lives six months under ground, and the other division of the year above ground, and emits its seed into the ball, and brings forth; and that there is not a female beetle. All then, in a word, who have spoken of divine things, both Barbarians and Greeks, have veiled the first principles of things, and delivered the truth in enigmas, and symbols, and allegories, and metaphors, and such like tropes. Such also are the oracles among the Greeks. And the Pythian Apollo is called Loxias. Also the maxims of those among the Greeks called wise men, in a few sayings indicate the unfolding of matter of considerable importance. Such certainly is that maxim, “Spare Time:” either because life is short, and we ought not to expend this time in vain; or, on the other hand, it bids you spare your personal expenses; so that, though you live many years, necessaries may not fail you. Similarly also the maxim “Know thyself” shows many things; both that thou art mortal, and that thou wast born a human being; and also that, in comparison with the other excellences of life, thou art of no account, because thou sayest that thou art rich or
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    renowned; or, onthe other hand, that, being rich or renowned, you are not honoured on account of your advantages alone. And it says, Know for what thou wert born, and whose image thou art; and what is thy essence, and what thy creation, and what thy relation to God, and the like. And the Spirit says by Isaiah the prophet, “I will give thee treasures, hidden, dark.”[746] Now wisdom, hard to hunt, is the treasures of God and unfailing riches. But those, taught in theology by those prophets, the poets, philosophize much by way of a hidden sense. I mean Orpheus, Linus, Musæus, Homer, and Hesiod, and those in this fashion wise. The persuasive style of poetry is for them a veil for the many. Dreams and signs are all more or less obscure to men, not from jealousy (for it were wrong to conceive of God as subject to passions), but in order that research, introducing to the understanding of enigmas, may haste to the discovery of truth. Thus Sophocles the tragic poet somewhere says: “And God I know to be such an one, Ever the revealer of enigmas to the wise, But to the perverse bad, although a teacher in few words,”— putting bad instead of simple. Expressly then respecting all our Scripture, as if spoken in a parable, it is written in the Psalms, “Hear, O my people, my law: incline your ear to the words of my mouth. I will open my mouth in parables, I will utter my problems from the beginning.”[747] Similarly speaks the noble apostle to the following effect: “Howbeit we speak wisdom among those that are perfect; yet not the wisdom of this world, nor of the princes of this world, that come to nought. But we speak the wisdom of God hidden in a mystery; which none of the princes of this world knew. For had they known it, they would not have crucified the Lord of glory.”[748] The philosophers did not exert themselves in contemning the appearance of the Lord. It therefore follows that it is the opinion of the wise among the Jews which the apostle inveighs against it. Wherefore he adds, “But we preach, as it is written, what eye hath not seen, and ear hath not heard, and hath not entered into the
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    heart of man,what God hath prepared for them that love Him. For God hath revealed it to us by the Spirit. For the Spirit searcheth all things, even the deep things of God.”[749] For he recognises the spiritual man and the gnostic as the disciple of the Holy Spirit dispensed by God, which is the mind of Christ. “But the natural man receiveth not the things of the Spirit, for they are foolishness to him.”[750] Now the apostle, in contradistinction to gnostic perfection, calls the common faith the foundation, and sometimes milk, writing on this wise: “Brethren, I could not speak to you as to spiritual, but as to carnal, to babes in Christ. I have fed you with milk, not with meat: for ye were not able. Neither yet are ye now able. For ye are yet carnal: for whereas there is among you envy and strife, are ye not carnal, and walk as men?”[751] Which things are the choice of those men who are sinners. But those who abstain from these things give their thoughts to divine things, and partake of gnostic food. “According to the grace,” it is said, “given to me as a wise master builder, I have laid the foundation. And another buildeth on it gold and silver, precious stones.”[752] Such is the gnostic superstructure on the foundation of faith in Christ Jesus. But “the stubble, and the wood, and the hay,” are the additions of heresies. “But the fire shall try every man’s work, of what sort it is.” In allusion to the gnostic edifice also in the Epistle to the Romans, he says, “For I desire to see you, that I may impart unto you a spiritual gift, that ye may be established.”[753] It was impossible that gifts of this sort could be written without disguise.
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    CHAPTER V. ON THESYMBOLS OF PYTHAGORAS. Now the Pythagorean symbols were connected with the Barbarian philosophy in the most recondite way. For instance, the Samian counsels “not to have a swallow in the house;” that is, not to receive a loquacious, whispering, garrulous man, who cannot contain what has been communicated to him. “For the swallow, and the turtle, and the sparrows of the field, know the times of their entrance,”[754] says the Scripture; and one ought never to dwell with trifles. And the turtle-dove murmuring shows the thankless slander of fault-finding, and is rightly expelled the house. “Don’t mutter against me, sitting by one in one place, another in another.”[755] The swallow too, which suggests the fable of Pandion, seeing it is right to detest the incidents reported of it, some of which we hear Tereus suffered, and some of which he inflicted. It pursues also the musical grasshoppers, whence he who is a persecutor of the word ought to be driven away. “By sceptre-bearing Here, whose eye surveys Olympus, I have a trusty closet for tongues,” says Poetry. Æschylus also says: “But I, too, have a key as a guard on my tongue.” Again Pythagoras commanded, “When the pot is lifted off the fire, not to leave its mark in the ashes, but to scatter them;” and “people on getting up from bed, to shake the bed-clothes.” For he intimated that it was necessary not only to efface the mark, but not to leave even a trace of anger; and that on its ceasing to boil, it was to be composed, and all memory of injury to be wiped out. “And let not the sun,” says the Scripture, “go down upon your wrath.”[756] And
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    he that said,“Thou shalt not desire,”[757] took away all memory of wrong; for wrath is found to be the impulse of concupiscence in a mild soul, especially seeking irrational revenge. In the same way “the bed is ordered to be shaken up,” so that there may be no recollection of effusion in sleep, or sleep in the day-time; nor, besides, of pleasure during the night. And he intimated that the vision of the dark ought to be dissipated speedily by the light of truth. “Be angry, and sin not,” says David, teaching us that we ought not to assent to the impression, and not to follow it up by action, and so confirm wrath. Again, “Don’t sail on land” is a Pythagorean saying, and shows that taxes and similar contracts, being troublesome and fluctuating, ought to be declined. Wherefore also the Word says that the tax- gatherers shall be saved with difficulty.[758] And again, “Don’t wear a ring, nor engrave on it the images of the gods,” enjoins Pythagoras; as Moses ages before enacted expressly, that neither a graven, nor molten, nor moulded, nor painted likeness should be made; so that we may not cleave to things of sense, but pass to intellectual objects: for familiarity with the sight disparages the reverence of what is divine; and to worship that which is immaterial by matter, is to dishonour it by sense. Wherefore the wisest of the Egyptian priests decided that the temple of Athene should be hypæthral, just as the Hebrews constructed the temple without an image. And some, in worshipping God, make a representation of heaven containing the stars; and so worship, although Scripture says, “Let us make man in our image and likeness.”[759] I think it worth while also to adduce the utterance of Eurysus the Pythagorean, which is as follows, who in his book On Fortune, having said that the “Creator, on making man, took Himself as an exemplar,” added, “And the body is like the other things, as being made of the same material, and fashioned by the best workman, who wrought it, taking Himself as the archetype.” And, in fine, Pythagoras and his followers, with Plato also, and most of the other philosophers, were best acquainted with the Lawgiver, as may
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