QUESTIONS OF

SCIENCE

QUESTIONS OF SCIENCE is an international series of books

published by Harvard University Press, Penguin Books, and
Editions Odile Jacob. Written by scientists of international
reputation, these books provide the nonspecialist reader
with an understanding of scientific thought at the frontiers
of our knowledge. The series will treat the enduring and
exciting questions in fields that particularly inform our sense
of place in the universe.
Dou 1 n House, Charles and Emm,1 Daru 1 in s home in Ke11t, iuhcrc
the ,(!eologist Clwrlcs Lyell met u 1 ith Dm·u 1 i11 in April 1856 ,wd
pcrs11,1dcd him that he shotild ptit ,111 ,1h.Hr,1a (f his 1 1 in,,s 011
cl'olution into pri11t.
 ONELONG
ARGUMENT
-------•-------

Charles Darwin and the Genesis of

Modern Evolutionary Thought

ERNST MAYR

HARVARD UNIVERSITY PRESS

QH371
.M336
1991

Copyright © 1991 by Ernst Mayr

All rights reserved
Printed in the United States of America
IO 9 8 7 6 ) 4 J 2

Design by Marianne Pcrbk

Library of Congress cataloging information is on bst p.1gc of book.

 For Michael T Ghiseli11
a11d Fra11k ]. Sul/0111ay,
u 1ho have contributed so ,nuch
to OllY ll11dersta11di11J! �r Danvi11
 Preface

A M o DERN Ev o Lu TI o NI s T turns to Darwin's work again

and again. This is not surprising, since the roots of all our
evolutionary thinking go back to Darwin. Our current contro­
versies very often have as their starting point so1ne vagueness in
Darwin's writings or a question Darwin was unable to answer
owing to the insufficient biological knowledge available in his
tin1e. But one returns to Darwin's original writings for more than
historical reasons. Darwin frequently understood things far more
clearly than both his supporters and his opponents, including
those of the present day.
An analysis of almost any scientific problen1 leads automati­
cally to a study of its history. The many unresolved issues in evo­
lutionary biology are no exception to this rule. To understand the
history of a scientific problem, however, one must appreciate not
only the state of factual knowledge but also the Zeitgeist of the
time. Any investigator's interpretation of his observations or ex­
periments depends mainly on this conceptual framework. For
many years a major objective of my historical studies has been to
discover the concepts-or sometin1es, even 1nore broadly, the
ideologies-on which the theorizing of certain historical figures
was based.
My interest in Darwin's thought arose in 111y university years,
but my more active preoccupation began in 1959 with the cente­
nary of the publication of On the OriJi11 o_f Species. I studied Dar­
win's writings even more intensely when I prepared an introduc­
tion to a facsimile of the first edition of the On��in, published in
1964. Curiously, most editions of this work available before that

..
• VI I •
 Preface

time were of the much-rev��ed sixth edition. The inexpensive

facsimile made wide distribUtion of the first edition of the On�<?in
possible for the first time since its original publication.
In the ensuing years I devoted myself to the study of Darwin's
work, and this effort culminated in The Growth o_f Biological
T houilzt: Diversity, Evolution, and Inheritance, published in 1982.
However, in this overview of the history of syste111atics, evolu­
tionary biology, and genetics it was impossible to present detailed
analyses of certain aspects of Darwin's work. These I treated in
separate lectures and papers, mostly presented at con1n1emora­
tive celebrations. Essays growing out of these lectures, along
with many other articles I had written on the history and philos­
ophy of biology, were collected in a volume entitled Toward a
New Philosophy o_fBiology: Obsen,ations ofan Evolutionist (1988).
While I was reviewing this rather technical and specialized col­
lection for a Gern1an translation, it occurred to n1e that a separate
volun1e devoted exclusively to Darwin and l)arwinisn1 might be
useful to students and lay people broadly interested in the role of
Darwin's thought in the history of ideas. As n1y starting point for
this new volume, I pulled from n1y collected essays the chapters
that were devoted principally to Darwin and Darwinism and be­
gan to edit them. Yet after judiciously pruning and shaping the
original eight essays, I realized that considerable gaps remained in
my presentation. I set about writing several new chapters, while
thoroughly reworking and rearranging material from the original
essays. The book that has resulted represents, I hope, a mature
reflection on Darwin's thought-one that emphasizes previously
neglected aspects of his work and clarifies controversial or con­
fused issues.
The last thirty years have been a period of unparalleled activity
in Dar\vin research, primarily owing to the discovery of the
��l)ar\vin papers''-notebooks, letters, unpublished manuscripts,
and so on. The first six volumes of Darwin's letters have now
been published, as has a single-volun1e edition of Darwin's note­
books. In addition to these prirnary docun1ents, books about
l)arwin and his life appear every fc\v years, and several of the111
arc very good. Yet even today 1nuch that is written about Dar\vin

• l'lll •
 Pnfacc

is si111ply wrong or, worse, n1alicious-in large part because the

author has failed to understand the concepts that underlie l)ar­
win's thought and its develop1nent, and the entrenched ideologies
that his "one long argun1ent" (a phrase he used in Chapter 14 to
describe the Or(gi11) was designed to oppose. This volun1e is an
attempt to correct son1e of those conceptual n1isunderstandings,
as well as to incorporate 111any of the latest findings of the still
exceedingly active l)ar\vin research program.
The facts of evolution as well as particular problems of phylo­
geny are hardly mentioned in this book. It is of little relevance
for evolutionary theory whether the ancestors of the n1olluscs
were metan1eric (segn1ented) or not (almost surely they were),
whether the coelenterates have the same ancestry as the flat­
worn1s, whether the tetrapods arose from lungfishes (as now
seen1s more probable) or coelacanths. An enormous literature al­
ready exists on these concrete proble1ns of phylogeny. Instead, I
have concentrated on the mechanisms of evolution and on the
historical development, from Darwin on, of the major concepts
and theories of evolutionary biology.
Finally, my attention to underlying concepts in this book rep­
resents an effort to combat a disturbing trend in our modern view
of basic scientific research. Science, in the minds of too many
scientists, is considered to be merely a sequence of discoveries or,
worse, a steppingstone to technological innovation. In my writ­
ings I hope to have established a better balance in the evaluation
of science by showing the close connection between the questions
raised within certain fields of science and other more general as­
pects of modern thought and inquiry. This endeavor automati­
cally leads to a consideration of Darwin, for no one has influenced
our modern worldview-both within and beyond science-to a
greater extent than has this extraordinary Victorian. We turn to
his work again and again, because as a bold and intelligent thinker
he raised some of the most profound questions about our origins
that have ever been asked, and as a devoted and innovative scien­
tist he provided brilliant, often world-shattering answers.

• lX •
. ... ,
 Contents

1 · Who Is Darwin? 1

2 · Confronting the Creationists: The First

Darwinian Revolution 12
3 · How Species Originate 26

4 · Ideological Opposition to Darwin's Five Theories 35

5 · The Struggle against Physicists and Philosophers 48
6 · Darwin's Path to the Theory of Natural Selection 68
7 · What Is Darwinis1n? 90
8 · A Hard Look at Soft Inheritance: Neo-Darwinism 108
9 · Geneticists and Naturalists Reach a Consensus:
The Second Darwinian Revolution 132
10 · New Frontiers in Evolutionary Biology 141

Ref crences 16 7

Glossary 177
Acknowledgments 189
Index 191

• Xl •
..
 Illustrations

Fnmtispiccc: Down House, Darwin's hotne in Kent. (Reproduced by

permission of Mary Evans Picture Library, London.)

Followin..� page 80:

J. S. Henslow (1796 -1861), Darwin's botany professor at Cambridge
University. (Reproduced by permission of the Syndics of the Fitzwil­
liam Museun1, Can1bridge.)
The H.M.S. Beagle, in the Straits of Magellan, 1833. (Reproduced by
permission of the National Maritime Museum, Greenwich.)
The route of the voyage of the Beagle, 1831-1836. (By Laszlo Mes­
zoly.)
The naturalist Jean Baptiste Lamarck (1744-1829). (Reproduced by
pern1ission of Bibliotheque centrale du Museutn National d'Histoire
Naturelle, Paris.)
Charles Lyell (1797-1875), geologist
The ornithologist John Gould (1804-1881). (Reproduced by permis­
sion of the Ipswich Borough Museutns and Galleries, Suffolk.)
En1ma Wedgwood Darwin (1808-1896) in 1840, Charles Darwin's
wife
Charles Darwin in 1840
The botanist Joseph Dalton Hooker (1817-1911 ), Darwin's friend and
supporter. (Reproduced by permission of The Royal Society, Lon­
don.)
The morphologist and paleontologist Richard ()wen ( 1804-1892),
who viciously attacked the Or(�in. (Reproduced by pcnnission of the
President and Council of the Royal College of Surgeons of England.)

• XI I I •
 II lustratio11s

Alfred Russel Wallace (1823-1913), co-discoverer with Dan-Vin of the

theory of evolution through natural selection
The morphologist, physiologist, and en1bryologist Thon1as Henry
Huxley (1825-1895), "Darwin's bulldog." (Reproduced by perrnission
of Lady Huxley.)
The botanist As a Gray ( 1 8 1 o- 1 8 8 8), Danv in's most i 111 portant sup­
porter in America
The Swiss-born naturalist Louis Agassiz (1809-1873). (Reproduced by
permission of the Museun1 of Con1parative Zoology. Harvard Uni­
versity.)
The biologist Ernst Haeckel (1834-1919), an enthusiastic supporter
and popularizer of Darwinism in Gerrnany. (Courtesy of the National
Library of Medicine, Bethesda, Maryland.)
August Weismann (1834-1914), the nineteenth century's greatest evo­
lutionist after Darwin. (Reproduced by pennission of the Museun1 of
Cornparative Zoology, Harvard University.)
Charles Darwin on the veranda at l)own House . (Reproduced by per­
n1ission of the New York Academy of Sciences . Courtesy of the Mu­
seun1 of Con1parative Zoology, Harvard University.)
ONE LONG ARGUMENT
..
 CHAPTER ONE
--------------•--------------

Who Is Darwin?

H I S TORI CA L PERI ODS arc do1ninatcd by distinct sets of

ideas which, taken together, fonn a well-defined Zeitgeist.
Greek philosophy, Christianity, Renaissance thought, the Scien­
tific Revolution, and the Enlightcn111ent arc exa111ples of sets of
ideas that don1inated their historical period. The changes fro111
one period to the next arc usually rather gradual; other changcs­
n1ore abrupt-are often referred to as revolutions. The n1ost far­
reaching of all these intellectual upheavals was the Darwinian rev­
olution. The worldview forn1cd by any thinking person in the
Western world after I 859, when 011 the Ort�{!ill o_f Sp ecies was pub­
lished, was by necessity quite different fron1 a world view forn1ed
prior to I 859. It is ahnost in1possible for a 111odern person to pro-
ject back to the early half of the nineteenth century and recon­
struct the thinking of this pre-Darwinian period, so great has
been the impact of Darwinisn1 on our views.
The intellectual revolution generated by 1)arwin went far be­
yond the confines of biology, causing the overthrow of son1e of
the most ba�ic beliefs of his age. For cxan1plc, Darwin refuted the
belief in the individual creation of each species, establishing in its
place the concept that all of life descended fron1 a con1n1on ances­
tor. By extension, he introduced the idea that hu111ans were not
the special products of creation but evolved according to prin­
ciples that operate every where else in the living world. Darwin
upset current notions of a perfectly designed, benign natural
world and substituted in their place the concept of a struggle for
survival. Victorian notions of progress and perfectibility were sc-

• 1 •
 l¥110 Is Darwin?

riously undermined by Dar.�in's demonstration that evolution

brings about change and aaaptation, but it does not necessarily
lead to progress, and it never leads to perfection.
Furthermore, Darwin established the basis for entirely new ap­
proaches in philosophy. At a time when the philosophy of science
was dominated by a methodology based on mathematical prin­
ciples, physical laws, and determinism, Darwin introduced the
concepts of probability, chance, and uniqueness into scientific
discourse. His work embodied the principle that observation and
the making of hypotheses are as in1portant to the advancement of
knowledge as experimentation.
Darwin would be remembered as an outstanding scientist even
if he had never written a word about evolution. Indeed, the evo­
lutionist J. B. S. Haldane went so far as to say that Darwin's most
original contribution to biology was not the theory of evolution
but his series of books on experin1ental botany published nearer
the end of his life (Haldane 1959:3 58). This achieven1ent is little
known an1ong nonbiologists, and the same is true for his equally
outstanding work on the adaptation of flowers and on anin1al
psychology, as well as his competent work on the barnacles and
his irnaginative work on earthworn1s. In all these areas Darwin
was a pioneer, and although in son1e areas n1ore than half a cen­
tury passed before others built on the foundations he laid, it is
no\v clear that Darwin attacked important problen1s with ex­
traordinary originality, thereby becon1ing the founder of several
now well-recognized separate disciplines (Ghiselin 1969). Darwin
\Vas the first person to work out a sound theory of classification,
one which is still adopted by the 1najority of taxonomists. His
approach to biogeography, in \vhich so n1uch en1phasis was
placed on the behavior and the ecology of organisms as factors of
distribution, is n1uch closer to n1odern biogeography than the
purely descriptive-geographical approach that don1inated bio­
geography for n1ore than a half-century after Darwin's death.
W ho was this extraordinary 1nan, and how did he con1e to his
ideas? Was it his training, his personality, his industry, or his ge-

• 2 •
 iVlw Is Da r1 1 1 i11 ?

nius that accounts for his success? Indeed, as \Ve shall sec, all were
involved.

The M an and His Work

Charles Darwin was born on February 1 2 , 1 809, at Shrewsbury,
England, the fifth of six children and the second son of l)r. Rob-
'--·

ert Darwin, an e111inently successful physician. His grandfather

was Erasmus Darwin, the author of Zoo11 0111 ia , a vvork which an­
ticipated his grandson's evolutionary interests by atten1pting to
explain organic life according to evolutionary principles. His
n1other, the daughter of Josiah Wedgwood, the celebrated potter,
died when Charles was only eight years old, and his elder sisters
tried to fill her place.
Fron1 his· earliest youth, l)arwin was a passionate lover of the
outdoors. As he hi1nself said, "I was born a naturalist." Every
aspect of nature intrigued hin1. He loved to collect, to fish and
hunt, and to read nature books. Shrewsbury was a country town
of about 2 0 , 000 inhabitants-a perfect place for the development
of a naturalist, 1nuch better than either a big city or a strictly rural
area would be.
School, consisting largely of the study of the classics, bored the
young naturalist intolerably. Before he turned seventeen years
old, Darwin's father sent him to the University of Edinburgh to
study medicine like his older brother. But n1edicine appalled
Charles, and he continued to devote much of his tin1e to the
study of nature. When it became clear that he did not want to
become a physician, his father sent him early in I 828 to Cam­
bridge to study theology. This seemed a reasonable choice, since
virtually all the naturalists in England at that time were ordained
ministers, as were the professors at Cambridge who taught bot­
any (J. S. Henslow) and geology (Adam Sedgwick). Darwin's
letters and biographical notes give the impression that in Cam­
bridge he devoted more time to collecting beetles, discussing bot­
any and geology with his professors, and hunting and riding with

. 3 .
 H'lzo Is Da rwin?

sin1 ilarly incl ined frien ds than to his p rescribed s tudies . Yet he did
I •

wel l in his exan1 inations , and when he took his B . A . in 1 8 3 1 he

s tood ten th on the l is t of nonhonors students . More i m portantly,
when Darwin had com pl eted his Cambridge years he was an ac­
con1 pli shed young naturalist .
l n1 n1ediatel y u pon finishing his stu dies Darwin recei ved an in­
vita tion to j oin H. M . S . Beagle as natura list and gen tlen1an co m­
panion of Captain Robert FitzRoy. Fitz Roy had been co n1 n1 i s­
s ioned to su rvey the co asts of Pa tagonia, Tierra del Fueg o , Chile,
and Pe ru to provide inforn1ation fo r 1naking better cha rts . The
voya g e wa s to be con1 p leted within two or three yea rs but actu­
ally las ted five . The Bea(�le left Plyn1outh on D cce 111 ber 2 7 , 1 8 3 1 ,
when Darwin was twen ty-t wo yea rs old , and retu rned to En g­
land on October 2 , 1 8 3 6 . l) arwin used these five years to their
fu llest exten t . In an erninentl y rea dable travel ogue (lounrnl (�f Re­
sea rch es) he tel ls about all the places he visited-volcanic and coral
islands , tropi cal fo rests in Brazi l , the vast pan1 pas of Patagonia , a
crossing of the Andes fro n1 Chile to Tucu1nan in Argentina , and
n1 uch , 1n uch 1nore . E very day brou ght unforgettable new ex pe­
rien ces , an invalua ble back ground fo r his life 's wo rk . He col lected
s peci1nens fro 1n widely differen t groups of organis 1ns, he dug
out i1n portant fossils in Pa tagonia , he devoted n1 uch of his tirne
to geolog y, but most of all he observed aspects of nature and
asked himsel f inn u 1nerable ques tions as to the how and why of
natural processes . He as ked H w h y " questions not only ab ou t geo­
log ical fea tu res and an imal life, but also about political and soc ial
situations . And it was his ability to ask profound q uestions and
his perseverance in trying to answer then1 that wo uld eventually
n1 ake Dar\vin a great s cientis t .
I n sp ite o f bein g des perately seasick every tin1e the ship en­
cou ntered rough wea ther, l)ar\vin n1 ana ged to read a grea t deal
of i 1n portan t scientific li tera tu re that he brought alon g on the
voyage. No scientific wo rk was n1 ore critical to his fu rther think­
i n g than the firs t two vo lu rnes of Charles Lyell 's Pri11 cip les (�( Ge­
olog y ( 1 8 3 2 ) , which not only gave l) ar\vin an advan ced course in
unifo nnitarian geolo gy-a theory that ch an ges in the ea rth 's sur-

. ., .
 Who Is Darw i11?

face h ave occu rred gra d u a l l y over lon g periods of ti n1e-but a l s o

introduced h i 111 to Jean Ba ptiste La 111 a rck 's a rgu1n ents fo r, and
Lyell 's a r g u 1n en ts a g a i n s t , evolutionary thinkin g .
When l) ar win boarded the Bea<'<lc h e still believed in the fi x ity
of s pec ies , a s did Lyell and all of his teachers at C a in bridge . Yet
during the S o u t h A 1n erican phase of the Beagle voyage l)ar win
n1 ade 1na n y o bs erva tions th a t grea tly puzzled hi111 and that shook
his belief in the fi x ity of s pecies . But it wa s really his visit to the
Galapagos i n S epten1 ber and () ctober 1 8 3 5 that prov ided hi111
with the cru cial evi dence, even though-being preoccupied dur­
ing his st ay with geological resea rches-he did not a t first real ize
i t . However, nine 111 on ths later, in J u l y 1 8 3 6 , he penned these
wo rds in his dia ry : " When I see these islands in sight of each other
and possessed o f but a scanty s tock of a n i 1nals , tenan ted by these
birds but slightly d i fferi n g in s t ructure and filling the s a 1n e place
in nature , I n1 ust s u s pect they arc varieties . . . if there is the
s l i gh test founda tion for these ren1 a rks , the zoo logy of the archi­
pelag oes w i l l be well wo rth exa1n inin g : for such fa cts would un­
der mine the s tabilit y o f s pecies " (B arlow 1 96 3 ) .
A fter his a rrival in E n g la n d in O ctober 1 8 3 6 Da rwin so rted his
collectio n s and sent them to various s pecialists to be des cribed in
the official a cco unt o f the Bea<(?le expedition . In M arch 1 8 3 7,
when the celebrated orn i thol ogis t John Gould insisted th a t the
m ockin g birds (Mi,n us) co llected b y Da rwin on th ree differen t is­
lands in the Galapagos were th ree dis tinct species rather than va­
rieties , as D a rwin had thou g h t , Dar win firs t understood the p ro­
cess of geo g raphic speciation ( S u lloway 1 9 8 2 ; 1 9 84) : that a new
s p ecies can develop when a population beco mes geographically
isola ted fro m its p a ren tal s pecies . Fu rthennore, if colonists de­
ri ved fro m a sin gle South A m erican ancestor cou l d beco n1 e th ree
s pecies in the Galapagos , th en a l l the s pecies of 1n ockingbirds on
the m ainland co uld h ave been derived from an ancestral s pecies ,
and so cou l d have , at an earlier time, the species of related gen era ,
and so fo rth . N u m e ro us s tatements in Darw in's writin gs con fi nn
that fro m the s p ring o f 1 8 3 7 on he fi nnly believed in the gradual
o rigin of n e w s pecies through geographic s pec iation , and in the

. 5 .
 Who Is Darwin ?

theory of evolution by common descent (see Chapter 2). But an­

other year and a half woulct pass before Darwin figured out the
mechanism of evolution, the principle of natural selection. This
happened on September 28, 1 83 8, as he was reading Malthus's
Essay on the Principle of Popu lation (see Chapter 6).
In January 1 839 Darwin married his cousin En1n1a Wedgwood ,
and in September 1 842 the young couple n1oved fron1 London to
a country house in the small village of Down (Kent), sixteen
miles south of London, where Darwin lived until he died on
April 19, 1 882. Darwin's health required the n1ove to a quiet place
in the country. After he had passed his thirtieth year there were
often long periods when he was unable to work more than two
or three hours a day-indeed, when he was co1nplctcly incapaci­
tated for rnonths on end. The exact nature of his illness is still
controversial, but all the syn1ptoms indicate a 111alfunctioning of
the autonon1ous nervous systen1 .

On the Origin of Species

Darwin did not publish his theories about evolution for another
t\venty years, even though he wrote some preli1ninary 1nanu­
script essays in 1 842 and 1 844. He devoted these years to his geo­
logical books and papers and to his n1onumental two-volume
n1onograph on the barnacles (Cirripedia) . Why did Darwin spend
eight years on this piece of taxonon1y instead of rushing into
print \vith his in1portant discovery of evolution by con1mon de­
scent through natural selection? Modern historical researches by
Ghiselin ( 1969) and others have clearly sho\vn that the barnacle
studies \Vere , for Darwin, an advanced graduate course in taxon-
01ny, 1norphology, and ontogenetic research and not at all a \vaste
of ti1ne. The experience he gained in these researches \vas an in­
valuable preparation for \vriting the Origin .
Finally , in April 1 85 6 , Dar\vin began to con1pose \vhat he con­
sidered to be his "big species book. " About t\vo years later, after
he had finished the first nine or ten chapters of this book , he re­
ceived a letter fro1n the naturalist Alfred Russel Wallace, \vho at

. 6 .
 Who Is Darwin?

that tin1e was collecting specimens in the Moluccas. This letter,

which Darwin received in June 1 85 8, was accompanied by a 111an­
uscript that Wallace asked Darwin to read and subn1it to son1e
journal if he found it acceptable. W hen l)arwin read the 111anu­
script, he was thunderstruck. Wallace had arrived at essentially
the same theory of evolution by con1mon descent throu gh natu­
ral selection as he . On J uly 1 , 1 85 8, Darwin's friends Charles Ly ell
and the botanist Joseph Hooker presented Wallace's n1anuscript,
together with excerpts fro1n Darwin's manuscripts and letters, at
a meeting of the Linnean Society of London. This presentation
amounted to a simultaneous publication of the findings of l)ar­
win and Wallace. Darwin q uickly abandoned his idea of finishing
his monumental work on the species and wrote instead \vhat he
called an "abstract," which beca1ne his famous On th e Or((?i11 o_f
Sp ecies , published November 2 4 , 1 8 5 9 .
The impact of the Origin was enormous. Quite rightly it has
been referred to as "the book that shook the world . " In its first
year the work sold 3 , 8 00 copies, and in Darwin's lifetime the Brit­
ish printings alone sold more than 2 7 , 000 copies. Several An1eri­
can printings, as well as innumerable translations, also appeared.
Nevertheless, only in our lifetime have historians understood
how fundamental the influence of this work has been. Every
modern discussion of man's future, the population explosion, the
struggle for existence, the pu rpose of man and the universe, and
man's place in nature rests on Darwin.
In the ensuing twenty -three years of his life Darwin worked
steadily on certain aspects of evolution he had not been able to
cover adequately in the Or(�in . In a two-volume work, Th e Va ri­
a tion o_f A n i,n a ls and Plan ts under Do m estica tion ( 1 86 8), he struggled
valiantly with the problem of how genetic variation originates.
In Th e Descen t o_f Ma n and Selection in Rela tion to Sex ( I 87 1) he
dealt with the evolution of the h uman species and expanded on
his theory of sexual selection. Tlze Exp ress ion o_f th e Emotions in
Ma n and A n im a ls ( 1 8 7 2 ) laid a foundation for the study of anin1al
behavior. Insecti vo rous Pla n ts ( 1 875) described the remarkable ad­
aptation of the sun-dew and other plants for catching and digest-

. 7 .
 Wh o Is Darwin?

ing insects . In Th e Effects of Cross- and Seif-Fertilization in th e Veg­

etable Kingdo,n ( 1 8 76) , in The Different Forms o_( Flou/ers 0 1 1 Plan ts o_(
th e Sa,ne Species ( I 8 77) , and in Th e Po wer o_f Mo vemen t in Plan ts
( 1 8 80) , D arwin discussed as pects of plant grow th and ph ysiol­
ogy, as indica ted in the titles . And finally in Th e Formation (�f Veg­
etable Mold, th rough the A ction of Worms , with Observations 0 1 1 Their
Habits ( 1 8 8 1 ) , he des cribed the i m portan t role pl ayed by earth­ •
wo rms in the fo rmation o f the topsoil .
How cou l d one man achieve so much in a lifetin1 e , pa rticularly
cons iderin g the cons train ts i 1nposed by his illness ? Only by re­
trea ting into the qu iet of the coun tryside, refusing to accept most
o ffered offices or mem berships on co m m ittees , and, through the
generosity o f his fa ther, living on his in herited in con1e was Dar­
win able to co n1 plete his tas k . Yet l) ar win was not a reclus e . He
kep t in to uch with the sc ien tific wo rld through an extensive cor­
res pondence and occa s ional visits to London , and he was a de­
vo ted husband and dedicated fa ther to his ten chi ldren .
Darwin \Vas described by his con ten1 p oraries as an extraordi­
n arily n1odes t , gen tle person who wen t out of his way to avoid
hurting anyone's feelin gs . He wo rked so hard because he had an
unq uenchable thirs t fo r learning , not in order to get adva ncement
or honors . In hi s publications he was a scientist's sc ien tist . He did
not w rite for the general public; when some o f his wo rks had
great popular su ccess, he was al ways astonis hed .
Ne vertheles s , D a rwin fought fo r the recogni tion of his find­
ings a n1ong s cien tists, and he was supported b y a sma ll band of
loyal friends-among them lyell, Hooker, and the morphologist
T. H. Hu xley, often referred to as Darw in's bulldog because i t
wa s he w h o in p u b l i c debates n1 ost often defen ded Dar\v in's theo­
ries . The 1nost fervent of D a r\vin's adn1 i rers \Vere the na turalis ts .
These included the codis coverer of evolu tion by natural selection ,
A . R . Wa llace, the en tomologist Henry Walter Ba tes , and the
naturalist Fritz Miiller.
To have a loyal group o f defenders \Vas very i m port ant because
l) a rwin was attacked with unusual ferocity. In 1 8 60 the Ha rva rd
University zoologist Louis A g assiz wrote that Darwin's theory
\Vas a " s cien tific n1istake, un true 1 11 its fa cts, uns cien ti fic in its

. 8 .
 Hl11 0 ls Da rwin ?

methods, and mischievous in its tendency. " The Or(�in was ex­
tensively reviewed in j ou rnals by the leading philosophers, theo­
logians, literary n1en, and scientists of the day. B y far the major­
ity of the revie ws were negative , if not extremely hostile (H ull
1 97 3 ) . Cu riously, this negative reception continued after Dar-
\Vin's death in I 8 8 2 and has lasted in certain circles to the pres­
ent day.

Darwin's Scientific Method

The years during which Darwin worked on the n1anuscript for
his big species book happened to be the same years in which the
field of study known as the philosophy of science originated in
England . While still a student, l)arwin had read with enth usiasm
John Herschel's Preli,ninary Discourse on the Study o_f Natural Philos­
ophy ( 1 8 3 0) , and this work continued to be one of his favorite
readings . He also read William Whewell's and John Stuart Mills'
books and tried conscientiously to follow their prescriptions for
the study of natural history (Ruse 1 97 5 6 ; Hodge 1 9 8 2) . This was
rather difficult, since the recommendations of the various authors
were often contradictory ; as a result, so were Darwin's own state­
ments on the subj ect. To satisfy some of his readers Darwin as­
serted that he followed "the true Baconian n1ethod" (Darwin
1 9 5 8 : I 1 9) , that is, straight induction. In reality, he "speculated"
on any subj ect he encountered. He realized that one cannot make
observations unless one has some hypothesis on the basis of
which to make the appropriate observations. Therefore, " I can
have no doubt that speculative men, with a cu rb on, make far the
best observers" (Darwin 1 98 8 : 3 1 7) . He stated his views most
clearly in a letter to Henry Fawcett. "About 3 0 years ago there
was much talk that geologists ought to observe and not to theo­
rize; and I well remember someone saying that at this rate a man
might as well go into a gravel pit and count the pebbles and de­
scribe the colors . How odd it is that anyone should not see that
all observation m ust be for or against some view if it is to be of
any service ! " (Darwin and Seward 1 90 3 ) .
Darwin's method was actually the time-honored method of the

. 9 .
 Mlho Is Darwin?

best natu ralists . They observe numerous p henomen a and al ways

try to understand the how and w h y of their observations . When
s om ething does not at once fall into place, they make a conj ecture
and test i t b y a dditional observations , leading either to a refuta­
tion o r stren gtheni n g o f the o ri ginal assum ption . This procedu re
does not fit well into the classical prescriptions o f the philosophy 'I

o f sc ience, because i t consists of continually going back and forth

between making observation s , posing questions , establishing h y­
potheses o r m o del s , testing them by making fu rther obs erva­
tions , and so forth . Darwin's speculation was a well-disciplined
p roces s , used b y h i m , as by every modern scien tis t, to give direc­
tion to the planning o f experi n1ents and to the collecting of fu r­
ther observations . I know of no forerunner of l)arwin who used
this method as consistently and with as n1u ch success .
That Darwin was a genius is hardly any longer questioned ,
som e o f his earlier detractors notwithstanding . B u t there n1 ust
have been a score of other biologists of eq ual in telli gen ce who
failed to match D arwin's achievemen t . What is i t that distin­
guis hes Darwin fro m all the oth ers ? Perh aps we can answer this
ques tion by inves tigating what kind o f scien tist Dar win was . As
he has said, he was fi rst and fore n1ost a naturalist . He wa s a s p len­
did observer, and like all other na turalists he was in teres ted in
organic diversity and in adap tation . Natu ralis ts are, on the
whole, describers and particularizers , but Darwin \Vas also a grea t
theoreti cian , so mething only very few naturalists have ever been .
I n that res pect Da rwin resembles much more some o f the leading
ph ysical sc ien tists of his day. But Darwin differed fro m the run­
of-the- n1ill natu ralists also in ano ther way. He \Vas not only an
obse rver but also a gifted and indefatiga ble experi men ter \v hen­
ever he dealt with a problen1 whose solution could be advanced
by an experi n1ent .
I think this s u ggests son1e o f the sou rces o f Dar \vin's greatnes s .
The universality o f his talen ts and in teres ts had prcadapted hin1
to beco n1e a bridge-bu ilder bet\veen fields . I t enabled hi n1 to use
his back g roun d as a natu ralist to theorize about so1ne of the n1ost
challenging problerns that pique our curios ity. And , in the face of

• 1 0 •
 Wh o Is Da ru 1 in ?

widespread beliefs to the contrary, Darwin was utterly bold in his

theorizing. A brilliant n1ind, great intellectual boldness, and an
ability to con1bine the best qualities of a naturalist-observer, phil­
osophical theoretician, and experimentalist-the world has so far
seen such a con1bination only once, and it was in the man Charles
Darwin.

• 1 1 •
 CHAPTER TWO
-------------•-------------

Confronting the Creationists :

The First Darwinian Revolution

A NE W D I SCO V ER Y I N SC I ENCE , such as the double-helix

structu re o f ON A, is usually accepted almost im mediatel y.
I f the assu med dis covery turns out to be based on an error or a
misinterpreta tio n , it quickly disappears fro n1 the literature . B y
con trast, resistance t o the introdu ction o f new theo ries , pa rticu­
larl y those that are based on new con cepts , is n1 uch s t ronger and
broader-based . Isaac New ton's theo ry that g ravitation is res pon­
sible fo r the n1 otion o f the planets requ ired some eigh t y years
before it was universally accep ted . A l fred Wegener's theory of
continental dri ft was publis hed in 1 9 1 2 but wa s generally adop ted
only fifty years la ter, a fter the acceptance of the theory of plate
tectonics .
Theo ries th a t either im plied or overtly assun1ed organic evolu­
tion had been p roposed fron1 B u tron ( 1 749) on , most expli citl y
by L a m a rck ( 1 809) , but were largel y i gnored or actively resisted .
When Charles D a rwin fi rst began to think abou t such problems ,
in his C a m b ridge days and on the Beagle, all of his teachers and
friends firmly believed that species do not change . They held this
belief in large part because of thei r religious views . The two
teachers in C a m b ridge to whom Da rwin was closes t , Hen slow
and Sed gwick , were both orthodox Christians and accepted the
dog1na of the Bible l iterall y, incl uding the story of creation . E ven
the geologist Ch arles Lyel l , whose wo rk pro foundly influenced
D a rwin's thinkin g-although Da rwin did not meet hi n1 person­
ally until a fter he retu rned from the Beagle voyage-was a theist
\vho believed that s pecies \Vere crea ted by God's hand . In all the

• 12 •
 Confron tin.._(;! the Cre ationists

wnttngs of the naturalists, geologists , and philosophers of the

period, God pl ayed a dominant role. They sa w nothing peculia r
in explaining otherwise puzzling phenomena as being caused by
God, and that included the question of how species originate .

The A r g u n1ent against Creationisn1

When Darwin decided in 1 8 27 to study for the tninistry, he too
was an orthodox Christian. As he said in his autobiography
( 1 9 5 8 : 5 7) , "As I did not then in the least doubt the strict and literal
truth of every word in the Bible ," he neither questioned the oc­
currence of miracles nor any other supernatural phenomena .
When he sailed on the Beagle , he reports, " I was quite orthodox ,
and I remember being heartily laughed at by several of the officers
for quoting the Bible as an unanswerable authority on some point
of morality " ( 1 9 5 8 : 8 5 ) . And yet it is evident that many of the
experiences he had during the five years of his voyage raised the
first dou bts in his mind about his religious beliefs. How could a
wise and good Creator permit the unspeakable cruelty and suffer­
ings of slavery? How could he instigate earthquakes and volcanic
eruptions that killed thousands or tens of thousands of innocent
people? Yet Darwin was far too busy with his work to become
obsessed by such disturbing thoughts. After his return from the
Beagle Darwin was more strongly influenced by his family's be­
liefs than by his Ca mbridge friends. As Ruse ( 1 979 : 1 8 1 ) has ob­
served: " His grandfather Erasmus was at best a weak deist, quite
able to believe in evolution . . . his father Robert, who had an
overwhelming influence on Darwin, was an unbeliever; his uncle
Josiah Wedgwood was a unitarian; and most important of all ,
Charles's older brother Erasmus had become an unbeliever by the
time Charles returned from the Beagle voyage."
But a more important influence on his changing beliefs than his
intellectu al environment was Darwin's own scientific findings .
Almost everything he learned in his natural-history studies was
more or less in conflict with Christian dogn1a . Every species had
numerous adaptations, from species-specific songs or courtships

• 13 .
 Co r�fronti11<� the Cre a tion ists

to specialized food and specifj.c enemies. According to the philos­

ophy of natural theology, which was widely accepted in England
at that time, God had designed and looked after all of these nun1-
berless details. They could not possibly be controlled by physical
laws, for these details were far too specific. Laws can control the
physical world, where adaptation was absent, but the specifi cities
and adaptations of the organic world required that God person­
ally look after every detail relating to thousands or, as we now
know, millions of species. Darwin could not accept this explana­
tion of the enormous diversity and adaptation he observed, and
he found himself more and more inclined toward natural 1necha­
n1sms.
Darwin's observations were also in conflict with the natural
theologians' belief in a perfect world. What the naturalist finds
instead are nu n1erous imperfections . How could all the species of
forn1er periods have become extinct if they had been perfect? As
Hull ( 1 973 : 1 26) has said rightly, "The god in1plied by . . . a real­
istic appraisal of the organic world was capricious, cruel, arbi­
trary, wasteful, careless, and totally unconcerned with the welfare
of his creations." Such considerations as these gradually drove
Darwin to the decision to try to explain the world \Vithout invok­
ing God or any kind of supernatural forces.
As \Ve will see in more detail below, Darwin adopted natural
selection in the fall of 1 8 3 8 , \vhich would suggest that he had
decided to reject supernatural explanations prior to that date. But
this conclusion has been vigorously denied by several authors,
whose views are seemingly well supported by Darwin's own
staten1ents in the Origi11 and elsewhere. There is now in the Dar­
win literature an extren1e spread of opinions, ranging from the
conclusion that Darwin was already an agnostic in 1 8 3 7 vvhen he
began to write his Notebooks, to the notion that he was still a theist
in 1 8 5 9 and became an agnostic only late in life. Ho\v could such
a \vide divergence of interpretation develop in vie\v of the n1as­
sive 111aterial relating to the issue in Darwin's O\vn notes, letters,
and publications? The answer is Darwin's own ambiguity. This
has been splendidly analyzed by Kohn ( 1 989) . Additional light on

• 1 4 .
 C01�fro11ti11J? tht' Creationists

cha n ges in D a r w in's thinking has been p rov ided by M o o re

( 1 9 8 9) . Readin g these accou nts (as well as ea rl ier ones by O s po­
va t , G reen e , G i l l e s p ie, M o o re , M a nier, and Ri cha rds) leads me to
the con clusion s that fo llow. However, Kohn has ri ghtl y said ,
every in terp reter of D a rwin's religiosity has ten ded to read into
D a rwin w h a t he wa n ted to fi n d , and presu mably I a n1 not es ca p­
i n g this wea kn ess either.
I t i s q u i te evident that p rior to t h e end ofJ u l y 1 8 3 8 Darwin had
m ade q u i te a few no tebook en tries that we re thoroughly " m ate r­
ialistic'' ( == agnos tic) . B u t J u l y 2 9 -3 1 , 1 8 3 8 , Da rwin visited the
Wed g woods at Maer in S t a ffo rdshire , and the co u rtship of
C h a rles and his cou s in E n1 m a began with tha t visit . E m m a was
an ortho d o x C h ristian a n d , as has o ften been pointed o u t , it be­
ca me clea r to D a rw i n that i t wo uld des troy thei r marria ge if he
was not ca u tious in the ex p ress ion of his reli gious views . M o re
than tha t , a s Kohn has said q u ite ri ghtly, " E mma beca me D a r­
win's m o del o f the co nven tional Vi ctorian reader" ( 1 9 8 9 : 2 2 6) . She
clearly h a d an effect on " t he contru ction o f D a rwin's texts . To me
this means one cru cial thin g , not a word of the a m b iguous God­
talk o f the Origin can be taken at fa ce va lue " (Kohn 1 9 8 9 : 2 26) . To
be s u re , it seen1 s t h a t D a r w in h i m self was still wavering . "A the­
is m b o th attra c ted and fri gh tened h i m " ( p . 2 2 7) . He was a ware of
the g reat u n k n own , and it wo uld have been a co mfo rt to him to
believe i n a s u p reme bein g . But a l l the p henomena of na ture he
enco un tered were con sis ten t with a straight fo rwa rd scien t i fi c ex­
plan a tion that d i d n o t invoke any su pernatu ral agen cies .
Thi rteen yea rs l a ter, in I 8 5 I , a n event occu rred in D a rwin's life
th a t t h o rou g h l y a ffected h i m . H e lost his beloved ten-yea r-ol d
d a u g h ter A n n i e , a child see m i n g l y perfect in h e r goodnes s . A s
Moore ( 1 9 8 9) describes , t h i s " c ruel " event s e e m s to have ex tin­
g u i s hed the l a s t tra ces o f thei s m in D a rwin .
Whether o n e wan ts to call h i m a dei s t , an agnostic, or an a the­
i s t , this m u ch is clear, that in the Origin D a rwin no longer re­
q u i red God as an exp lan atory fa ctor. C reation as des cri bed in the
B i b l e wa s contradic ted fo r D a rwin by almost every as pect o f the
nat ural wo rl d . Fu rtherm ore, c reation s i m p l y coul d not explain

• 15 .
 Confron ting th e Creatio n ists

the fossil record, nor the hierarchy in types of organisms that had
been proposed by the taxoriomist Carl Linnaeus, nor many of the
other findings of science. Yet almost all of Darwin's peers still
believed in some form of creation, and many of Darwin's con­
temporaries accepted Bishop Ussher's calculation that creation
had occurred as recently as 4004 years B . C .
By contrast, the geologists had long been aware of the im­
mense age of the earth, which would have allowed plenty of time
for abundant organic evolution. Another discovery of geology
that was most important and most disturbing for the creationist
was the discovery of abundant extinction. Already in the eigh­
teenth century the German naturalist Johann Friedrich Blumen­
bach and others had accepted ex tinction of formerly existing
types like an1n1onites, belemnites, and trilobites, and of entire
faunas, but it was not until Cuvier worked out the extinction of a
whole sequence of mammalian faunas in the Tertiary of the Paris
Basin that the acceptance of extinction became inevitable. The
ultimate proof for it was the discovery of fossil n1astodons and
mammoths, animals so huge that any living survivors could not
possibly have remained undiscovered in some remote part of the
globe.
Three explanations for ex tinction were offered. According to
Lamarck, no organism ever became extinct ; there simply was
such drastic transformation that formerly existing types had
changed beyond recognition. According to another school, one
to which Louis Agassiz belonged, each former fauna had become
extinct as a whole through some catastrophe and was replaced by
a newly created, more progressive fauna. This had happened, ac­
cording to Agassiz, fifty tin1es since the earth was formed. Such
catastrophism was unpalatable to Lyell, who produced a third
theory consistent with his uniformitarianism. He believed that
individual species becan1e extinct one by one as conditions
changed and that the gaps thus created in nature were filled by
the introduction of new species through some presun1ably super­
natural n1eans. Lyell's theory was an atten1pt at a reconciliation

• I 6 ·
 C01�fro11ti11(.{? th e Creation ists

between those who recognized a ch anging wo rld of long du ra­

tion and those who su pported the ten ets of crcationi s 111 .
The qu es t ion o f precisel y how these new s pec ies we re intro­
duced was left unanswered by Lyel l . He bequeathed this problen1
to Da rwin , who in due tin1e made it his n1 ost i 111 portan t resea rch
pro g ra m . D a rwin thus approached the proble1n of evolu tion in
an en tirel y d i fferen t m anner fro n1 La1narck . For Lan1arck , evolu­
tion wa s a s t ri ctly vertical phenon1enon, proceed ing in a single
dimensio n , that of tin1 e . Evolution for him was a 111 ove mcnt
fro m less perfect to 111 ore perfect , fro1n the n1 ost p ri111 itive in fu­
sorians u p to the n1 a n1 n1 als and 111 an . Lama rck 's Ph i l osop h ie Zoo l ­
og iqu e was the p a radi g 1n of ve rtical evolutionis1n . S pecies pl ayed
no role in L a 1n a rck 's thinkin g . New s pecies origina ted all the
time by s p o n taneous generation fro 1n inanin1a te n1a tter, but this
p roduced only the sin1 plest infusorians . Each newly establis hed
evo l ut ionary line g radually moved u p to ever greater perfection ,
as organi s m s a d a p ted to their environment and passed along to
their o ffs p ri n g t hese newly acquired traits .
D a rwin was unable to bu ild on this foundation but rather
started fro m the funda men tal question that Lyell had bequeathed
to hi m , n a mel y, how do new species originate? A l though Lyell
had a p pealed t o " intermediate causes " as the source of the new
species , the p ro cess was nevertheless a fo rm of s pecial creation .
" Spec ies may h ave been crea ted in su ccession at such times and at
such p la ces as to enable them to n1 ultiply and endure for an a p­
pointed period and occu py an a p pointed space on the globe"
(Lyell I 8 3 5 , 3 : 99 - 1 00) . For Lyell , each creation was a carefully
planned even t . The reason why Lyell , like Henslow, Sedgwick,
and all the o the rs o f D a r win's s cien tifi c frien ds and corres pon­
den ts in the m iddle of the 1 8 3 0s , accep ted the una l terable con­
stancy o f s pecies was ulti matel y a philoso phical one. The con­
stancy of s pecies - th at i s , the inability of a species , once crea ted ,
to change- wa s the one piece o f the old dogma of a created wo rl d
that rem ained invi olate a fter the concepts o f the recen cy and con­
stancy of the p h ys ical world had been abandoned .

• 1 7 ·
 Confronting th e Crea tion ists

Evidence for the Gradual Evolution and

Multiplication of Species
No genuine and testable theory of evolution could develop until
the possibility was recognized that species have the capacity to
change, to become transformed into new species, and to multiply
into several species. For Darwin to accept this possibility re­
quired a fundamental break with Lyell's thinking. The question
which we must ask ourselves is how Darwin was able to emanci­
pate himself from Lyell's thinking, and what observations or con­
ceptual changes permitted Darwin to adopt the theory of a trans­
forming capacity of species.
As Darwin tells us in his autobiography, he encountered n1any
phenomena during his visit to South America on the BeaJ:le that
any modern biologist would unhesitatingly explain as clear evi­
dence for evolution. Yet even when sorting his collections on the
homeward voyage, and realizing that the "varieties" he observed
" would undermine the stability of species," Darwin at that date
(approximately July 1 8 3 6) had not yet consciously abandoned the
concept of constant species (Barlow 1 963 ) . This Darwin appar­
ently did in two stages. The discovery of a second, sn1aller species
of rhea (South American ostrich) led him to the theory that an
existing species could give rise to a new species, by a sudden leap
or saltation. Such an origin of new species had been postulated
scores of times before, from the Greeks to Robinet and Mauper­
tuis (Osborn 1 894) . Sudden new origins, however, are not evo­
lution. The diagnostic criterion of evolutionary transformation is
gradualness.
The concept of gradualism, the second step in Darwin's con­
version, was apparently first adopted by Darwin when the orni­
thologist John Gould, who prepared the scientific report on Dar­
win's bird collections, pointed out to him that there \Vere three
different enden1ic species of 1nockingbirds on three different is­
lands in the Galapagos. Darwin had though t they ,vere onl y va­
rieties (Sulloway I 9 826) . The 1nockingbird episode ,vas of partic­
ular i1nportance to Darvvin for t,vo reasons . The Galapagos

• 18 .
 Confrontin�{! th e Creation ists

enden1 ics were quite si,nilar to a species of n1 ockingbirds on the

South A n1erican 111 ainland and clea rl y derived fro n1 it . Th us, the
Galapa gos birds were not the result of a single saltation , as l) ar­
win had postu lated for the new s pecies of rhea in Pa tagonia, but
had gradually evolved into th ree separate but si 1nilar species on
th ree different islands . This fa ct helped to convert Da rwin to the
con cept of gradual evolution (see Cha pter 3 ) . Even n1ore i 111 por­
tan t was the fa ct that these th ree different species had branched
off fro m a single paren tal species , the mainland mockin gbird­
an observation that gave Da rwin a solution to the problen1 of
speciation -tha t is , how and why species n1 ultiply.
For a beli ever in saltational evolution , specia tion meant the
sud den change of a species into a different one . For a believer in
transformational evolution , speciation n1 eant the gradual chan ge
of a species into a differen t one in a phyletic lineage. But neither
of these two theories explained the origin of the enormous or­
gan ic diversity Darwin saw a round hin1 . There are at least ten
million species of ani1nals and aln1ost two million species of
plants on the Earth today, not to n1 ention the countless kinds of
fungi, p rotis ts, a n d p roka ryotes . Even though in Darwin's day
onl y a fra ction of this number was known , the question why
there are so many s pecies and how they originated was already
being asked. Lamarck had ignored the possibility of a multipli­
cation of species in his Ph ilosop h ie Zoologique ( 1 809) . For hi m , di­
versity was produced by differences in rates of adaptation, and
new evolutionary lines originated by spon taneous genera tion, he
though t . In Lyell 's steady-state world, species number was con­
stant, a n d new species were introduced to replace those that had
become extin ct . Any thou ght of the splitting of a species into
several daughter species was absent among these earlier authors .
A solution to the problem o f species diversi fication required an
en tirely new approach , and only the naturalists were in the posi­
tion to fi n d i t . Leopold von Buch in the Canary Islands , Darwin
in the Galapagos, Moritz Wa gner in No rth Africa , and A . R .
Wallace in A m azonia an d the Malay A rchipelago were the pio­
neers in this endeavor. They each discovered nu merous popula-

• 19 .
 Confronting the Creation ists

tions that were i n a l l conceivable

' ...
in termedia te stages of species
fo rmatio n . The s h a rp discontinuity between species that had so
i m p ressed John Ray, C a rl Linnaeus , and others was now called
into question b y a con tinuity a m ong species .
The p ro b lem o f how these new species and in cipient sp ecies
co m e in to bei n g wa s cl ari fied for Darwin by the Galapag os
mocking b i rds . These speci mens showed that new s pecies can
originate b y what we now call geographical (or allopa tric) specia­
tion . This theory o f speciation says that new s pecies n1 ay ori gi­
n a te by the gradual genetic transformation of isolated popula­
ti ons . These isolated popul ations beco me in the cou rse of time
geo g ra p h i c races or subs pecies , and Dar win reali zed that they
may beco me new s pec ies , if isolated suffi ciently lon g . B y this
thought Da rwin founded a branch of evolu tion is n1 which , fo r
short, we rn ight des i g n a te as horizontal evolu tion i s n1 , in con tra s t
wi th t h e s t rictly vertical evolut ionism of Lama rck . T h e two
terms deal with two en tirely different as pects of evolutio n , even
though the processes res ponsible fo r these as pects p roceed si mul­
taneously. Ve rtical evolutionism deals wi th adaptive changes in
the t i m e di mens ion , while ho rizontal evolution i s n1 deals with the
origin o f new diversity in the space di mension , that is , with the
origin of inc ipient s pecies and new species as populations move
into new envi ron mental ni ches . These s pec ies en rich the diversity
of the organic wo rld and are the potential fo un ders o f new higher
taxa and of new evolu tionary dep artu res that will occupy new
adap tive zones .
Fron1 1 8 3 7 on , when Da rwin first reco gnized and solved the
problem o f the o rig in of di versity, this vertical /h orizontal duality
of the evolutionary p rocess has been with us . Unfortunately, only
a fe w authors have had the breadth of thought and experience of
D a rwin that wou ld a l low them to deal si multaneou sly with both
as pects o f evolu tion . Instea d , paleontologists and geneti cists have
con centrated on ve rtical evol ution , while the maj o rity of na tural­
ists have s tud ied the origin of diversity as reflected in the horizon­
tal p rocess of the n1 ti l tiplica tion of s pecies and the origin of
highe r taxa .

• 20 •
 C01�fron ti11g th e Creation ists

For Da rwin , horizontal thinking about spcciation pcnnitted

the solu tion of th ree i1n portant evolutionary problems : ( 1 ) why
and how species n1ultiply; (2) why there a rc discon tinui ties be­
tween n1 aj or groups o f organisn1 s in nature, when the con cept of
gradual evolution would seem to i m pl y countless subtle grada­
tions between all g rou ps; and (3 ) how higher taxa could evolve .
But perhaps the mos t decisive con sequence o f the discovery of
geo gra phic s peciation wa s that it led Darwin auton1atical l y to a
branching concept of evol u tion . This is why branching en tered
Darwin's notebooks at such an early stage (sec below) .
Despite these insights of Da rwin and the n aturalists, the deep
significance of the concept of speci ation has been cla rified only
very slowly. As we will see in n1orc detail in Chapter 3 , in the
decades a fter the publica tion of the Orig in o.f Species Darwin him­
self vacillated in his understanding of how species n1 ultipl y, and
he con tinued to s t ru ggle with problcn1s surrounding the origin
of diversity fo r the res t of his life. Nevertheless , his realization
that any evolution a ry theory 1nust son1ehow account for the
multiplication of s pecies was a pillar of Darwin's evolutionary
thought from I 8 3 7 onward .

The Theory of Com mon Descent

The case of the s pecies o f Galapagos 1nockingbirds provided Dar­
win with an addition al i m portant new insight . The th ree species
had clearly descended fro m a sing le ancestral species on the South
American con tinen t . Fro m here it was only a small step to pos­
tulate that all mockingbirds were derived from a co n1mon ances­
tor - indeed , that every grou p of organisms des cended fron1 an
ances tral s pecies . This is Darwin's theory of con1 mon descen t. B y
con trast, for those w h o accep ted the concept o f the sea /a naturae
(scale of n atu re, or Great Chain of Bein g) -and in the eighteen th
cen tury this included most naturalists to a lesser or greater ex­
ten t - all o rganisms were part of a single linear scale of ever­
growing perfection . La marck still adhered , in prin ciple, to this
concept even thou gh he allowed for some branching in his classi-

• 21 •
 Confronting th e Crea tion ists

fication of the major phyla . . Peter Simon Pallas and others had
also published branching diagrams, but it required the categorical
rejection of the sea/a naturae by Cuvier in the first and second dec­
ades of the nineteenth century before the need for a new way to
represent organic diversity became crucial .
A group known as quinarians experimented with indicating re­
lationship by osculating circles, but their diagrams did not fit re­
ality at all well . The archetypes of Richard Owen and the Natur­
p h ilosop hen strengthened the recognition of discrete groups in
nature, but their use of the term "affinity" in relation to these
groups remained meaningless prior to the acceptance of the
theory of evolution. For Agassiz and Henri Milne-Edwards,
branching reflected a divergence in ontogeny, so that the adult
forms were far more different than the earlier e1nbryonic stages .
Fron1 all these exampies it is evident that the static branching dia­
grams of nonevolutionists are no n1ore indications of evolution­
ary thinking than branching flow charts in business or branching
diagrams in administrative hierarchies.
Apparently very soon after Darwin understood that a single
species of South A n1erican n1ockingbird had given rise to three
dau ghter species in the Galapagos Islands, he seemed to realize
that such a process of multiplication of species, combined with
their continuing divergence, could in due time give rise to differ­
ent genera and still higher categories. The members of a higher
taxon then would be united by descent from a comn1on ancestor.
The best way to represent such a con1mon descent would be a
branching diagram. Already in the summer of 1 8 3 7 Darwin
clearly stated that "organized beings represent an irregularly
branched tree" (Notebook B : 2 1 ) , and he drew several tree dia­
gran1s in which he even distinguished living and extinct species
by different symbols.
By the time Darwin wrote the Origin , the theory of common
descent had become the backbone of his evolutionary theory, not
surprisingly so because it had extraordinary explanatory powers.
Indeed , the manifestations of comn1on descent as revealed by
con1parative anatomy, co1nparative e1nbryolog y. systernatics (in

· 22 ·
 Co1�fronti11i the Creation ists

its sea rch for " the n a tu ral s ys te111 " ) , and biogeography beca 1n e
t h e n1 ain evidence fo r the occurren ce o f evolution i n the years
after I 8 5 9 . Reci procally, these biological disci plines , which up to
I 8 59 had been p rin1 a ril y des cri ptive , now beca me causal sc ien ces ,
with co 1n m o n descen t providing an explanation fo r nearly every­
thing that h a d p reviously been puzzlin g .
The con cept o f co n1 n1on des cen t was not en ti rely ori ginal with
D a rwin , however. B u ffon had a l ready cons idered it fo r close rel­
atives s u ch as horses and a sses ; but not accepting evolu tion , he
had not ex ten ded this thou ght s ys tematically. There a re occa­
sional s u g ges tions of co 1n m on descen t in a n u n1 ber of other p re­
D a rwinian w ri ters , but his torians so fa r have not n1 ade a careful
search for earl y a dherents o f co n1 111 on ancest ry. The theory was
definitel y n o t u pheld by Lan1 a rck , who, although he proposed
the occasional s pli tting of " m asses " (hi gher ta xa) , never thou gh t
in terms o f a s plittin g o f s pecies and regular branching . For him
descent was l inear des cen t wi thin each phyletic line, and the con­
cep t of co m m o n descen t was a lien to hi m .
The theory o f co m m on descen t , once proposed, is so s i m ple
and so obvious that i t is hard to believe Darwin was the first to
have a do p ted i t consistently. I ts i m p ortance was not only that i t
had s u ch g reat explanatory powers but a l s o t h a t i t provided a
unity for the living world that had been previously missin g . Up
to 1 8 5 9 people had been i m p ressed primari l y by the enormous
divers ity o f life, fro m the lowes t plan ts to the highes t verteb ra tes ,
but this diversity took on an entirely different co mp lexion w hen
it was rea l ized that i t all could be traced back to a com m on origin .
The fi n al proof o f this , o f course, wa s not sup pl ied until our time,
when i t was demons trated that even ba cteria have the same ge­
netic code as animals a n d plan ts .
None o f D a rwin's theories was accep ted as enthusiastical l y a s
co m m on descen t . E verything that had seemed to b e arbitra ry o r
chaotic in n a t u ral histo ry u p t o tha t point n ow began to make
sense. The a rchet y p es o f O wen and of the co mparative ana to­
mists cou l d n ow be ex p lained as the heritage fro n1 a com m on
an ces tor. The en tire Linnaean hierarch y suddenly beca me q u ite

. 23 .
 Ccn�fron ting the Cre a tion ists

logical, because it was now apparent that each higher taxon con­
sisted of the descendants ofa still more remote ancestor. Patterns
of distribution that previously had seemed capricious could now
be explained in terms of the dispersal of ancestors. Virtually all
the proofs for evolution listed by Darwin in the Origin actually
consist of evidence for common descent. To establish the line of
descent of isolated or aberrant types became the most popular
research program of the post-Origin period, and has largely re­
mained the research program of comparative anato1nists and pa­
leontologists almost up to the present day. To shed light on co1n­
n1on ancestors also became the program of comparative
embryology. Even those who did not believe that ontogeny re­
capitulates phylogeny often discovered sin1ilarities in e1nbryos
that were obliterated in the adults. These sin1ilaritics, such as the
chorda in tunicatcs and vertebrates, or the gill arches in fishes and
terrestrial tetrapods, had been totally n1 ystifying until they were
interpreted as vestiges of a con1n1on past. Though there are still a
nun1ber of connections an1ong higher taxa to be established, par­
ticularly among the phyla of plants and invertebrates, there is
probably no biologist left today who would question that all or­
ganisn1s now found on the earth have descended from a single
origin of life. This Darwin anticipated when he suggested that
"all our plants and animals [have descended] fro1n some one
forn1, into which life was first breathed" ( 1 859:484).
But perhaps the n1ost i1nportant consequence of the theory of
con1n1on descent was the change in the position of n1an. For
theologians and philosophers alike, n1an was a creature apart
from the rest of life. Aristotle, Descartes, and Kant agreed in this,
no rnatter how n1uch they disagreed in other aspects of their phi­
losophies. Darwin, in the Origin , confined hin1self to the cau­
tiously cryptic ren1ark, "Light will be th ro\vn on the origin of
Man and his history " (p. 48 8). But Ernst Haeckel, T. H. Huxley,
and in 187 1 Darwin hi1nself den1onstrated conclusively that hu-
1nans n1ust have evolved from an ape-like ancestor, thus putting
then1 right into the phylogenetic tree of the ani1nal kingdon1.
This \Vas the end of the traditional anth ropoccntrisrn of the Bible
 Co,�fron ti11._1? th e Creatiomsts

and o f the philoso phers . This application o f the theo ry of co111-

mon des cen t to h u 1na n s , however, encoun tered vigorous o p po­
sition . To j udge fro1n conten1 porary cartoons , n o Da rwinian idea
wa s less accep table to the Victorians tha n the deri vation of n1 an
from a p ri 111 ate an ces tor. Yet today this derivation is not only re­
n1 arkably well s u bs tan tia ted by the fossil reco rd , bu t the bio­
che1nical and chro 1n o s o 1n a l sin1il arity of n1 an an d the A frican
apes is so g reat that i t is puzzling why they are so rel atively differ­
ent in morphology and brain devel o p n1 en t . The prin1 a te o rigin
of man, as fi rs t s ugges ted by Darwin , in1 mediately raised ques­
tions about the o rigin of 1n ind and cons ciousness that a re con tro­
vers ial to this d ay.

The Fate o f D arwin's First Revolution

Darwi n's theory o f evol ution a s such (together with the nortEetl­
stancy o f s pecies) , his theory o f co m mon des cen t , and his theory
that s pecies multi pl y were victorious within a rem arkably s ho rt
time. The victory of these three theories is the first Darwinian
revolu tion . Within fi fteen years o f the pu blication of the Origin
hardly a qualified biologist was left who had not beco me an evo­
l utionis t . The evolu tionary origin of n1 an was s till una ccep table
to some with rel i g ious com mitments but was taken as firmly es­
tablis hed by the an thro pological p rofess ion .
E ven though a theory o f the m u l ti plication of s pecies is now
taken fo r g ra nted as an essen ti a l co m ponent of evolutionary
theory, i t is s till con trovers i a l how this multiplication co mes
about . That much, if not m os t , s peciation is geographical is gen­
erally accep ted , but what o ther forms of s pecia tion m ay also ex­
ist, a n d how frequen t they a re, is s till an unsettled p roblem , as we
will see in the next cha p ter.

. 25 .
 C HAP T E R T H R E E
--------------•-------------

How Species Originate

F RoM THE s u M M E R o F 1 8 3 7 o N , Darwin collected notes on

the great book he was going to write, and he referred to it in
his notes and in his correspondence always as "the species book. "
When finally published in 1 8 5 9 , it was called by hin1 011 tlze Origin
of Species . Darwin was fully conscious of the fact that the change
from one species into another one was the most fundamental
problem of evolution. Indeed, evolution was, almost by defini­
tion, a change from one species into another one. The belief in
constant, unchangeable species was the fortress of antievolution­
ism to be stormed and destroyed. Once this was accomplished, it
would not be difficult to adduce other evidence in favor of evo­
lution.
In view of this central position of the problem of species and
speciation in Darwin's life work, one would expect to find in the
Origin a satisfactory and indeed authoritative treatment of the
subject. This, curiously, one does not find. Indeed, the longer
Darwin struggled with these concepts, the more he seemed to
becon1e confused. In the end, the Origin was a superb treatment
of the theory of common descent and a great plea for the efficacy
of natural selection, but it was vague and contradictory both on
the nature of species and the mode of speciation.

Wha t Is a S pecies ?
After his crucial conversation \Vith John Gould about the Gala­
pagos 1nockingbirds in 183 7, Dar\vin continued to struggle \vith

• 26 .
 Ho w Species Originate

the problen1 of how to define a species; but for that matter, so did
virtually all other naturalists during the ensuing 1 5 0 years. It will
help to understand l)arwin's problem if I give a short overview of
the four n1ajor species concepts that developed during this pe­
riod.

TH E TY POLOGICAL SPECI ES CONCE PT

A typological species is an entity that differs fron1 other species
by constant diagnostic characteristics. This was the species con­
cept of Linnaeus and Lyell and was supported by those philoso­
phers from Plato to modern ti n1es who consider species to be
" natural kinds" or "classes. " The concept was entirely consistent
with the belief in creationis111 ("that which was separately created
by God") and with the philosophy of essentialism ("that which
has an essence of its own" ; see Chapter 4) . However, this species
concept has three major practical weaknesses, which is the reason
why it has been largely given up in recent times. First, it forces
its adherents to consider as species even different variants within
a population. Second, the invalidity of this concept is demon­
strated by the high frequency of so-called sibling species-spe­
cies which are indistinguishable on the basis of their appearance
but which do not interbreed in nature. These species cannot be
discriminated under the typological species concept. And third,
it forces us to recognize as full species many local populations that
differ by one diagnostic character from other populations of the
species.

TH E NOMINALIST SPECI ES CONCE PT

According to this concept, only individual objects exist in nature.
Similar objects or organisms are bracketed together by a name,
and this subj ective action of the classifier determines which ob-
jects arc combined into one species. Species, therefore, are
merely arbitrary mental constructs. That species have no reality
in nature has been the claim of the nominalists from the Middle
Ages right up to the writings of some recent philosophers. By
contrast, the real ity of species has been supported consistently by

• 27 .
 Ho w Sp ecies Orig inate

naturalists up to the p resent . day. There is no more devasta tin g

I ..

refuta tion of the nominalistic claim than the fact that primitive
natives in New Guinea , with a Stone A ge culture, recognize as
s pecies exact ly the same entities of natu re as Weste rn taxo no­
mists . I f species were something purel y arbit rary, it would be to­
ta ll y i m p robable for rep resentatives of two drastically different
cultures to arri ve at iden tical s pecies deli mitations .

T H E EV O L UTIO N A R Y S PE C I ES C O N C EPT
Paleontolo gists w h o study species distribu ted i n the ti1ne dimen­
sion have always been rather dissatisfied with a species defini tion
derived fro m the nond imensional s pecies con cep t of the local nat­
u ralis t . What paleon tol ogists were looking fo r wa s a species con­
cept that would be particularl y suitable fo r the discri 1nination o f
fossil species . This need eventually led the naturalist G . G . Simp­
son ( 1 96 1 : 1 5 3 ) to this s pecies definition : "A n evolutionary s pecies
is a lineage (an ances tral-descen dent seq uen ce of populations)
evolving separately fro n1 others and with its own unitary evolu­
tionary role and tendencies . " Wiley ( 1 9 80, 1 9 8 1 ) and Will mann
( 1 9 8 5 ) have proposed slightl y modified versions of Simpson's def-
1n1t1on .
A n evolutionary definition o f species has not been widely ac­
cepted because it is on l y applicable to monotypic s pecies , since all
isola tes that " evol ve separately " would also have to be recognized
as s pecies . Fu rthermore , how is one to describe and determine
the " unitary evolutionary role and tendencies " of a popul ation or
tax on ? The main obj ective of the evolutionary species definition
wa s to permit a clear deli mitation of a species in the tin1e dimen­
sion , but this hope turned out to be illu sory in all cases of gradual
s pecies transforn1a tion . The exact determination of the origin of
a new s pecies can be n1 ade only in cases of insta ntaneous s pecia­
tion (as in polyploid y) , and the exact endpoint of a species can
only be detern1 ined in the case of extinction . The biolo gical spe­
cies definition , a s we will see, is equall y quali fied to determine
these t wo points , and has other ad vantages going fo r it as well .

• 28 .
 Ho w Species Or((?inate

THE BIOLOGI CAL SPECIES CONCEPT

This s pecies concept is based on the ob servation of local n a tu ral­
i s ts that at a given locality, populations of differen t s pec ies coexist
b u t d o n o t in terb reed with each other. This I articula ted in the
defini tion : " S pecies a re g rou ps o f interb reed ing natu ral popula­
tions that a re rep roductively isolated fro m other such g rou ps . "
S t a te m en ts i n D a rwin's notebooks show that b y I 8 3 7 h e had
g i ven u p the typologi cal s pecies con cept and had devel oped a s pe­
cies con cept based on rep rodu cti ve isolation . For in stance , " M y
defini tion o f s pecies has n o thing to do with h y b ridity, is si m pl y
an ins tin ctive i n1 pulse to keep separa te, which will no doubt b e
overco m e [ o r else n o h ybrids wo uld ever b e produced ] , b u t until
i t is, these ani n1 a l s a re distinct sp ecies " (C : 1 6 1 ) . There are re­
peated referen ces in the notebooks to n1 utual " repugnance " of
s pecies t o i n tercrossin g . " The dislike of two species to each other
is evidently an i n s tinct; and this prevents breedin g " ( B : 1 97) . " D ef­
ini tion o f s pecies : one that re1nains at large with constant cha rac­
ters , together w i t h other bein gs o f very near structure" ( B : 2 1 3 ) .
Fo r D arwin , s pecies s ta tus had little if anything to do with de­
gree of d i fference : " Hence species n1 ay be good ones and differ
sca rcely in any ex tern al character " ( B : 2 1 3 ) .
D arwin a d he red to this biological s pecies concept for about the
next fi fteen yea rs . But then h e beca n1e rather confused , particu­
larl y a fter h e tried to apply his zoological findings to p lants . A s
we s h all s ee , h e con side red the variety (which i n animals i s a sub­
s pecies ) to be an inci pien t s pecies , and he encoun tered no diffi cul­
ties with this concept as long as he dealt with ani1nals . Howeve r,
when throu g h Hooker a n d other botanical frien ds D a rwin ca n1 e
t o s tu d y varieties of plants , he d i d n o t rea lize that t h e bo tanica l
ter m i n ol o g y w a s rather differen t fro n1 t h a t of zoology. P l a n t va­
rieties very o ften were individual va riants within a local popu la­
tion , a n d to consider them incip ient s pecies not only caused
problems fo r the explanation o f s pec iation but also fo r the delim­
itation of s pecies against va rieties , and of s pecies against one an­
other. There were a n u m ber o f other develo p 1nen ts in these years

. 29 .
 Ho w Sp ecies Origin a te

which induced Darwin to B,ive up his biological species concept

and return to a more or less typological species definition (Sullo­
way 1979). The twenty-five pages on species and speciation in his
unfinished big book manuscript (Darwin 1975 : 250 -274) contain
so many contradictions that they are almost embarrassing to
read.
The species concept at which Darwin finally arrived is clearly
described in the Origin . There is nothing left of the biological
criteria of the notebooks, and his characterization of the species
now is a mixture of the typological and nominalist species defi­
n1t1ons.

Systema tists . . . will not be incessantly haun ted by the shadowy

doub t whether this or that form be in essence a species. This I feel
sure . . . will be no slight relief . . . Syste111a tists \vill have only to
decide . . . whether any forn1 be sufficien tly constant and d istinct
from other fonns, to be ca pable of definition ; and , if defina ble ,
whether the differences b e sufficiently in1 portant t o deserve a spe­
cific nan1e . . . The only distinction between species and well­
marked varieties is , that the latter are known, or believed , to be
connected a t the presen t day by intermediate gra da tions . . . In
short , we sha l l have to take species in the san1e manner as those
n a tura lists trea t genera , who admit that genera are n1erely artificial
combinations made for convenience . This may not be a cheering
prospect ; but we shall a t least be freed from the v ain search for the
und iscovered and undiscoverable essence of the term species "
( I 8 5 9 : 4 84-4 8 5 ) ·

The example set by Darwin was followed by just about every

taxonon1ist and evolutionist in the nineteenth century except for
a few enlightened field naturalists. It clearly was the species con­
cept of the Mendelians. The counter movement in the t\ventieth
century, initiated by Jordan, Poulton, Stresen1ann, and other pro­
gressive workers, ultimately resulted in the biological species
concept's being widely accepted.
However, each of the three n1ajor species concepts (typologi­
cal, evolutionary, and biological) has a certain legitimacy in some

. JO .
 Ho w Species Or(gi11ate

areas of biological research , even today. Which of them one

adopts may depend on the type of research one does . The mu­
seu m taxonomist, as well as the s tratigrapher, may find the ty po­
logical s pecies concept n1ost useful, never mind how clea rl y it is
refuted by the existen ce of sibling s pecies and strikingl y differen t
variants (phena) . B u t anyone working with living populations ,
restricted to one place and time, finds any species con cept other
than the biological one to be unsatis factory. Fin ally, the paleon­
tologis t , part of whose endeavor it is to delimit fossil species taxa
in the vertical sequence of stra ta, cannot help but pay attention to
the time dimensio n .
Modern biologists are almost unanimously agreed that there
are real discontinuities in organic nature, which delimit natural
entities that a re designated as s pecies . Therefore, the species is
one of the basic foundations of almost all biological disciplines .
Each species has different biological cha racteristics , and the anal­
ysis and com p arison of these differences is a prerequisite fo r all
other research in ecology, behavioral biology, co1nparative mor­
phology and physiology, molecular biology, and indeed all
branches of biology. Whether he realizes it or not , every biologist
works with species .

What Is Speciation?
Paleontologists , conditioned t o vertical thinking , considered spe­
ciation to be the change of a phyletic lineage over time. But since
there is a steady _e xtinction of species , where do the new species
come fro m ? This has been the p roblem from Lamarck and Lyell
on . The ans wer which Darwin found was that species not only
evolve in time but also multiply. Speciation , since Darwin, has
come to mean the production of new, reproductively isolated
populations .
That the m ultiplication of species was a n1aj or evolutionary
phenomenon became clear to Darwin as soon as he followed up
the conseq uences of his new understanding of the taxono m y of

. 31 •
 Ho w Sp ecies Originate

the Galapago s mockin gbirds . . But it was really not until the co m­
ing of the new s ys tematics .. and the evolutionary synthesis (see
Chapter 9) that speciation became the focal point of evolutionary
research at the s pecies level .
Considering Darwin's uncert ain ties about the natu re of spe­
cies , it is perhaps not surp rising that he va cillated about the mode
o f s peciation . I t must be re membered that at first Darwin be­
l ieved in Lyell 's sudden , s a ltational " introduction " of new spe­
cies . I t was John Gould 's pronouncement in M arch 1 8 3 7 that each
of the three popul ations of mockingbirds in the Gala pagos Islands
was a sepa rate s pecies that started Darwin along an entirely new
path of thinking about the origin of diversity. Obviously, the dif­
feren t m ockin gbirds on the three islands in the Galapagos had all
descended from colonists of the South A merican mainl and spe­
cies ; but the three popula tions had evolved in a slightly different
n1 anner on ea ch o f the three islands . This led Darwin to a dopt the
theory of s peciation by the gradual n1 odification of popula tion s
that were geo graphically isolated fro m their paren t s pecies . And
this was D a rwin's theory of s pecia tion until the early 1 8 5 0 s . I t was
not a new theory, because a similar one had previou s l y been pro­
posed by von Buch ( 1 8 2 5 ) . La ter, Wa gner ( 1 8 4 1 ) and Wallace
( 1 8 5 5 ) also in dependently arrived at the same conclusion .
Darwin considered isolation on islands as the principal specia­
tion n1echani s m , and he seems to have had difficul ties in expl ain­
ing s pecia tion on continen ts . At one time, to account for the rich
species di versity in South A frica , he pos tulated large-scale geo­
logical changes -up and down movements of the crus t-during
which South A frica was temp orarily converted into an archi­
pela go, setting the s tage fo r abundant geographical specia tion .
Just a s Darwin's botanical researches induced hin1 to change his
s pecies concept, so did these resea rches , at least in part , induce
Da rwin to accep t , in addi tion to geographic speciation , a process
\Ve 110\v call s y rnpatric speciation. The exact reasoning by which
Da rwin arrived at this ne\v explanation is s till not fu lly explored
( S ul lo\vay 1 9 79 ; M ayr 1 9 8 2 ; Kohn 1 9 8 5 ) . O\ving to the prevailin g
ty pological thinking , geographic races of animals \Vere called va-

. 32 .
 How Species Or(g in ate

rieties in Darwin's tin1e. In plants, however, as n1entioned above,

the tern1 variety had a dual n1eaning . I t was applied not only to
geographic races (subspecies) but also to individual variants
(phena) within a single population. Darwin, on the basis of his
zoological studies, had concluded that varieties of animals were
incipient species. When he transferred this concept of the role of
varieties in speciation to plants, he was induced to consider also
individual variants within a population as incipient species. Dar­
win concluded, therefore, that in addition to geographic specia­
tion there is also a process of sympatric speciation, which he de­
fined as the origin of a new species by ecological specialization
within the range of the parental species. The survival and flour­
ishing of the new sympatric species, it was said, was made pos­
sible by its shift into a new niche, thus removing it from compe­
tition with the parental species. Natural selection would lead to
an ever g reater difference (character divergence) of the two com­
peting new species. Darwin developed this concept of sympatric
speciation by character divergence in the years I 8 54 to 1 8 5 8
(Kohn 1 9 85) and continued to support it even though once in a
while he would concede that maybe geographic isolation was
nearly always necessary.
I t was this claim of sympatric speciation that involved Darwin
in a bitter controversy with the explorer and naturalist Moritz
Wagner ( 1 86 8, 1 8 8 9), who insisted that geographic isolation was
absolutely indispensible for speciation. Unfortunately Wagner
beclouded the issue by also insisting that natural selection could
not operate unless the population was isolated. What we now
know about speciation indicates that Wagner 's arguments on the
whole were more valid than Darwin's. Nevertheless, Darwin's in­
sistence on the frequency of sympatric speciation p revailed, and
p rior to the evolutionary synthesis sympatric speciation was con­
sidered a frequent and, as far as the entomological literature is
concerned, perhaps the prevailing mode of speciation.
In his early notebook statements on species (B: 1 97, 2 1 3 ;
C: 1 6 1 ), Darwin clearly saw species as reproductively isolated en­
tities. By the 1 8 50s, however, his attention concentrated on spe-

. 33 .
 Ho w Sp ecies Originate

cies as adapted entities. He would describe how they became

adapted to a new niche, but he failed to account for their repro­
ductive isolation from the parental species. His reasoning was ag­
gravated by a return from populational to typological thinking
(see Chapter 4) . " If a variety were to flourish so as to exceed in
number the parent species, it would then rank as the species and
the species as the variety, or it might come to supplant and exter­
minate the parent species, or both might coexist, and both rank
as independent species" ( 1 8 5 9 : 5 2). How could the new variety
coexist as an independent species unless it had acquired reproduc­
tive isolation? This question is particularly troublesome, given
that Darwin, in an argument about the origin of isolating mech­
anisms among species, definitely rejected the capacity of natural
selection to establish a reproductive barrier (Mayr 1 9 596; Sullo­
way 1 979).
Looking over all of Darwin's writings on species and specia­
tion, one cannot escape the impression that he was rather con­
fused about the issues and that he not infrequently contradicted
himself. A major reason for his confusion can be traced to his
puzzle1nent over the origin of genetic variation. The clear state­
ments about mechanisms of speciation that we now can make
n1ore than 1 2 5 years later are based on our understanding (as far
as it goes) of genetics. Despite his failure to reach a definitive
conclusion about species and speciation, Darwin deserves credit
for having pointed out the problems and for having clearly stated
various alternative solutions.

. 34 •
 C H APT E R F O U R
--------------•--------------

Ideological Opposition to
Darwin's Five Theories

I N BoT H s cHo LA RLY and popular literature one frequently

finds references to "Darwin's theory of evolution," as though
it were a unitary entity. In reality, Darwin's "theory " of evolution
was a whole bundle of theories, and it is impossible to discuss
Darwin's evolutionary thought constructively if one does not dis­
tinguish its various components. The current literature can easily
leave one perplexed over the disagreements and outright contra­
dictions among Darwin specialists, until one realizes that to a
large extent these differences of opinion are due to a failure of
some of these students of Darwin to appreciate the complexity of
his paradigm.
The term "Darwinism," as we will see in Chapter 7, has nu­
merous meanings depending on who has used the term and at
what period. A better understanding of the meaning of this term
is only one reason to call attention to the composite nature of
Darwin's evolutionary thought. Another reason is that one can­
not answer the question correctly of how and when "Darwin­
ism" was accepted in different countries throughout the world
unless one focuses on the various Darwinian theories separately.
What Darwin presented in 1 859 in the Origin was a compound
theory, whose five subtheories had very different fates in the
eighty years after Darwin.
One particularly cogent reason why Darwinism cannot be a
single monolithic theory is that organic evolution consists of two
essentially independent processes, as we have seen: transforma­
tion in time and diversification in ecological and geographical

. 35 .
 Ideological Opp osition to Da rwin 's Five Theories

space. The two processes ��quire a minimum of two entirely in­

dependent and very different theories. That writers on Darwin
have nevertheless almost invariably spoken of the combination of
these various theories as " Darwin's theory" in the singular is in
part Darwin's own doing. He not only referred to the theory of
evolution by common descent as "my theory," but he also called
the theory of evolution by natural selection "my theory," as if
common descent and natural selection were a single theory.
Discrimination among his various theories has not been helped
by the fact that Darwin treated speciation under natural selection
in chapter 4 of the Or(gin and that he ascribed many phenon1ena,
particularly those of geographic distribution, to natural selection
when they were really the consequences of co1nmon descent.
Under the circumstances I consider it necessary to dissect Dar­
win's conceptual fran1ework of evolution into a number of major
theories that formed the basis of his evolutionary thinking. For
the sake of convenience I have partitioned Darwin's evolutionary
paradigm into five theories, but of course others n1ight prefer a
different division. The selected theories are by no means all of
Darwin's evolutionary theories; others were, for instance, sexual
selection, pangenesis, effect of use and disuse, and character di­
vergence. However, when later authors referred to Darwin's
theory they invariably had a combination of some of the follow­
ing five theories in n1ind:

( 1 ) Evolu tion as such . This is the theo ry that the wo rld is not constant
nor recen tly crea ted nor perpetually cycling b u t rather is steadily
ch anging and that organisms are transforn1ed in time .
(2) Common descent . This is the theory that every g roup o f organis ms
des cen ded fro m a co m mon ancestor and that all g roups of orga­
nis m s , including animal s , plants , and n1 icroorganis m s , ul ti-
1natel y go back to a single origin of life on earth .
( 3 ) A1u ltip lica tion of sp ecies . This theory explains the origin of the
enorn1ous o rganic diversity. It postulates that s pecies n1 ultiply,
either b y s plitting into daughter species or by u budding ," that is ,
b y the establish men t of geog raphically isolated fo under popu la­
tions that evolve into new species .

. 36 .
 Ideologica l Opposition to Darwin 's Five Th eories

(4) Gradualism . According to this theo ry, evolu tionary chan ge ta kes
place th rou gh the g radual change of populations and not by the
sudden (saltational) p roduction of new in dividuals th at represent
a new type .
( 5 ) Natural selection . According to th is theory, evolutionary change
co mes about through the abundant production of genetic va ria­
tion in every genera tion . The relatively few in dividuals who sur­
vive, ow in g to a particu larly well-adapted co rn bina tion of inher­
i table ch ara cters , give rise to the next generation .
For D a rw in h i m self these fi ve theories were a p p a rently a unity,
and so meone might cla i n1 that in deed these five theo ries a rc a
l o gically ins eparable p a ckage and that Da rwin was q uite correct
in t rea tin g the1n a s s u ch . This clai 1n , howeve r, is re fu ted by the
fact , as den1ons trated in Ta ble 1 , that most evolutionists in the
im mediate p o s t- I 8 59 period-that is, authors who had accep ted
the fi rs t t heo ry- rej ected one or several of D a rwin's other fou r
theories . This s hows tha t the fi ve theories a re not one indivisible
who l e .
There a re several reasons why D a rwin's five theo ries h a d such
differen t fates . One reason i s that abundant evidence was alread y
ava ilable to s u p p o rt some o f them . This was true, fo r example,

TABLE I
The composition of the evolutionary theories of various evolutionists.
A l l these authors accepted a fifth component , that of evolution as
opposed to a constant, unchanging world.

Com mon Mul tiplication Gradu- Natural

descent of species alism selection

Lamarck No No Yes No
Darwin Yes Yes Yes Yes
Haeckel Yes ;> Yes In part
Neo-
Lamarckians Yes Yes Yes No
T. H. Huxley Yes No No ( No) d
de Vries Yes No No No
T. H. Morgan Yes No (No) d Unin1 portant

a. Parentheses indicate ambivalence or contradiction .

. J7 .
 Ideologica l Opp osition to Da rwin 's Five Th eories

fo r the theory o f co m mon. descen t . I n the case of the theory of

natu ral selection , however, the eviden ce for or again st \Vas c-on­
tradictory, and it required generations of research time befo re the
con troversial poin ts could be settled . A more im portant reason ,
however, for the del ay in the acceptance of some of Darwin's
theories was that they were in conflict with certain ideologies
dominant at D ar win's tim e .
D u ring t h e th ree centuries preceding the pub lication o f the
Origin , Europe had been in the throes of a continuous intellectual
u pheava l , punctua ted b y the S cientific Revolution of the six­
teen th and seventeenth cen turies and the Enlighten ment of the
eighteen th cent ury. Why did it take so long fo r evolution to be
seriously proposed ? And why did Darwinism face such an uphill
battle a fter it wa s proposed? The reason is that Darwin challenged
so me of the basic beliefs of his age. Four of then1 we re pillars of
C h ristian dog m a :
( 1 ) A belief in a constant wo rld . In spite of Lam arck and the Natu rp h i­
losop h en , it was still widely, i f no t almost universally, accep ted in
1 8 5 9 that except fo r minor perturba tions (floods , volcanis m ,
mountain building ) the wo rld had n o t changed materially since
creation . And in spite of Buffon , Kant, Hutton , Lyel l , and the
ice age theory, the prevailing opinion was still that the wo rl d had
been crea ted ra ther recently.
(2) A belief in a created wo rld . Species and other taxa were believed to
be unchanging, and therefore the existing diversity of the l iving
wo rld could only be due to an act of crea tion . This was a single
crea tion as believed by the orthodox C hri stians or repea ted cre­
ations , either of whole biota as believed by the so-cal led progres­
sionists (for exa mple, Agassiz) , or o f individual species as pro­
posed by Lyell .
( 3 ) A belief in a wo rld des(gned by a u 1 ise and ben ({?n Creator. Even
though the world had its im perfections , it ,vas the bes t of the
possible worlds , according to Leibniz . The adap ta tion of orga­
nis n1s to their physical and li ving environn1ent \Vas perfect be­
cause it had been designed by an on1nipo ten t Crea tor.
(4) A belief in th e un ique position o_f man in th e creati011 . The ,vo rld ,vas
anthropocentric in the eyes of the Chris tian rel igion as \vell as in

. J8 .
 Ideolo�'<ica l Opp osition to Da rwin 's Five Theories

the view of the foremost philosophers . Man had a soul , some­

thing anin1als d id not have . There was no possible transition
fron1 anin1al to man .

I n addition to these four religious beliefs , there were th ree sec­

ular philosophies that were also in conflict wi th certain o f Dar­
win's theo ries . These secular beliefs were :
( 5 ) A belie_{ in th e ph ilosophy of essentia l ism (see below) .
( 6) A be/ ie_f in an in terp reta tion (�f th e causa l p rocesses of natu re as th ey had
been elaborated by th e p h ysicists (see Chapter 5) .
(7) A bel ie_f in ')11zal ca uses " or teleolo�'<Y (see Chapter 5 ) .
The theories of evolu tion proposed by Darwin challenged all
seven of these traditional and well-entrenched views , though not
every one of D arwin's five theories was in conflict with all of
them .

External Factors in Scientific Revolutions

For the last sixty years there has been a good deal of controversy
in the history of science over the question whether scientific rev­
olutions or, indeed, any conceptual changes in science a re due to
dis coveries and new observations made within the field (" inter­
nalism " ) or are the result o f outside influences ( " externalism " ) .
The Marxist thesis , fi rs t promoted by Hessen ( 1 9 3 1 ) , that the so­
cioeconomic mil ieu is decisive in initiating conceptual changes in
science was particularl y actively debated during part of this pe­
riod .
Researches du ring this period have made it clear that one must
distinguish between two sets of external factors : socioeconomic
and ideological ones . It has also become quite clear that these two
sets of factors differ funda men tall y in their impact on s cien ce .
Socioeconomic factors , it would seem , have very little influence
on the acceptance or rej ection of scien tific ideas , as was shown by
M ay r ( 1 98 2 : 4- 5 ) and b y many other historian s . I f the industrial
revolution and the socioeconomic situation had been res ponsible

. 39 .
 Ideo logica l Opp os ition to Darwin 's Five Th eories

. ..
for the theory of natural selection, this theory should have been
most palatable to the British public of the mid-nineteenth cen-
tury. But this was not the case at all. On the contrary, the theory
of natural selection, as we shall see, was almost unanimously re-
jected by Darwin's contemporaries. It was evidently not a reflec­
tion of the socioeconon1ic situation.
The other set of external factors-ideological ones-by con­
trast, had a powerful effect on the acceptance of Darwin's theo­
ries. What is remarkable and rarely sufficiently emphasized is that
it was not the prevailing ideologies almost universally adopted in
Britain (and much of the rest of the world) in the first half of the
nineteenth century that inspired Darwin and gave him his new
theories. On the contrary, the prevailing Zeitgeist was utterly op­
posed to Darwin's thought and prevented a universal acceptance
of so1ne of his new ideas for more than a hundred years. Indeed,
as shown by the many recently published books and papers that
still atten1pt to refute Darwinisn1, many Darwinian ideas are still
not yet full accepted, owing to the continuing power of the op­
posing ideologies.

Fro m Essentialis m to Popula tion Thinking

Of the seven ideologies challenged by Darwin's theories, none
was rnore deeply entrenched than the philosophy of essentialism.
Essentialism had dominated Western thinking for more than 2000
years, going back to the geometric thinking of the Pythagoreans.
They pointed out that a triangle, regardless of the combination of
angles, always has the form of a triangle. It is discontinuously
different from a quadrangle or any other kind of polygon. The
triangle is one of the limited nun1ber of possible forn1s of a poly­
gon. In an analogous 1nanner, all the variable phenomena of na­
ture, according to this thinking, are a reflection of a limited num­
ber of constant and sharply delimited eide or essences.
Essentialism, as a definite philosophy, is usually credited to Plato,
even though he was not as dogmatic about it as son1e of his later
followers, for instance the Thon1ists. Plato's cave allegory of the
 Ideologica l Opposition to Darwin �, Fi ,,c Theories

world is well known: W hat we see of the phcno1nena of the world

corresponds to the shadows of the real objects cast on the cave
wall by a fire. We can never sec the real essences. Variation is the
manifestation of in1perfect reflections of the underlying constant
essences.
All of Darwin's teachers and friends were, n1ore or less, essen­
tialists. For Lyell, all nature consisted of constant types, each cre­
ated at a definite tin1e. " There arc fixed lin1its beyond which the
descendants fro1n con1n1on parents can never deviate fro111 a cer­
tain type." And he added e111phatically : " It is idle . . . to dispute
about the abstract possibility of the conversion of one species into
another, when there are known causes, so much n1orc active in
their nature, which 1nust always intervene and prevent the actual
accomplishment of such conversions" (Lyell 1 83 5 : 1 62). For an es­
sentialist there can be no evolution: there can only be a sudden
origin of a new essence by a major n1utation or saltation.
Virtually all philosophers up to Darwin's tin1e were essential­
ists. W hether they were realists or idealists, n1aterialists or non1i­
nalists, they all saw species of organisms with the eyes of an es­
sentialist. They considered species as "natural kinds," defined by
constant characteristics and sharply separated from one another
by bridgeless gaps. The essentialist philosopher William Whewell
stated categorically, " Species have a real existence in nature, and
a transition from one to another does not exist" ( 1 840, 3 :626). For
John Stuart Mill, species of organisms are natural kinds, just as
inanimate objects are, and " kinds are classes between which there
is an impassable barrier."
Essentialism's influence was great in part because its principle
is anchored in our language, in our use of a single noun in the
singular to designate highly variable phenomena of our environ­
ment, such as mountain, home, water, horse, or honesty. Even
though there is great variety in kinds of n1ountain and kinds of
home, and even though the kinds do not stand in direct relation
to one another (as do the members of a species), the sin1ple noun
defines the class of objects. Essentialistic thinking has been highly
successful, indeed absolutely necessary, in mathematics, physics,

. 41 •
 Ideolog ica l Opp osition to Da rwin 's Fi i1 e Theories

and logic. The observati0n of nature seemed to give powerful

support to the essentialists' claims. Wherever one looked, one
saw discontinuities-between species, between genera, between
orders and all higher taxa. Such gaps as between birds and mam­
mals, or beetles and butterflies, were mentioned often by Dar-
. ' . .
win s cnttcs.
In daily life we largely proceed essentialistically (ty pologically)
and become aware of variation only when we compare individu­
als. He who speaks of "the Prussian," "the Jew," "the intellectual"
reveals essentialistic thinking. Such language ignores the fact that
every human is unique; no other individual is identical to him.
It was Darwin's genius to see that this uniqueness of each indi­
vidual is not limited to the human species but is equally true for
every sexually rep roducing species of animal and plant. Indeed,
the discovery of the importance of the individual becan1c the cor­
nerstone of Darwin's theory of natural selection. It eventually re­
sulted in the replacement of esscntialism by population thinking,
which emphasized the uniqueness of the individual and the criti­
cal role of individuality in evolution. Darwin no longer asked, as
had Agassiz, Lyell, and the philosophers, "What is good for the
species?" but "What is good for the individual?" (Ghiselin 1969).
And variation, which had been irrelevant and accidental for the
essentialist, now became one of the crucial phenomena of living
nature.

Fro m Saltationis m to Gradu alism

Many of Darwin's contemporaries who accepted the fact of evo­
lution nevertheless were incapable of population thinking, owing
to their ideological commitment to essentialism. As we have
seen, they accepted instead a concept of evolution based on the
sudden production of new species through saltations. Saltational
evolution is a necessary consequence of essentialisn1: if one be­
lieves in evolution and in constant ty pes, only the sudden produc­
tion of a ne\v ty pe can lead to evolutionary change. That such
saltations can occur and indeed that their occurrence is a necessitv
 Ideologica l Opposition to Darwin 's Fi11e Th eories

are old beliefs . A l n1ost all theories of evolution described b y Os­

born in his histo ry o f evolution , From th e Greeks to Da rwin ( 1 894) ,
were saltational theories , that is , theories of the sudden origin of
new kinds . A fter the publica tion of the Orig in , many biologists
who accepted evolution as such (Darwin's first theory) neverthe­
less, because they were essentialists , turned to s altational theories
to explain the proces s of evolution .
One can distinguish three kinds of sal tationist theories : ( 1 ) ex­
tinct s pecies a re replaced by newly created ones that arc 1nore or
less at the same level as those tha t they replace (Lyell 1 8 3 0 - 1 8 3 3 ) ;
(2) extinct species are replaced by new crea tions at a higher level
of organization (progressionis ts, such as William Buckland ,
Sedgwick , H u g h Miller, A gassiz) ; (3 ) new species originate
through saltations of pre-existing species (E . Geo ffroy Saint­
Hilaire , Da rwin in Patagonia, Galton , Goldschmidt) .
Saltational evolu tion is best called " transmutation ," because the
production of new s pecies or new types is discontinuous , owing
to the sudden creation of a new essence . But transmutation
should not be con fu sed with another concept of evolution known
as transformational evolution . According to this concept, evolu­
tion consists of the gradual transformation of a thing from one
condition to another. Lamarck 's concept of evolution was tra ns­
formational, designating a co mpletely gradual process , a chan ge
due to a tren d toward perfection or adj ustmen t to the environ­
men t . To the bes t of my knowledge Lamarck was the first author
to propose a consistent theo ry of gradual transformation . After
1 800 , but before 1 8 5 9 , the idea of g radual evolution was accepted
by a considerable n u m ber of authors on the Continent, but in a
vague manner and unsup ported by adequate evidence .
As Lewontin ( 1 9 8 3 ) has pointed ou t, Da rwin , by contrast, in­
troduced a new concept of evolution that was entirely different
from saltational evolu tion or transformational evolution . Ac­
cording to D arwin's concept, which we can designate as " varia­
tional evolution , " varia tions are produced in every generation ,
an d evolution takes pl ace because only a small number of va riants
survive to rep roduce . No longer is a con crete object transformed ,

. 43 .
 Ideolog ica l Opp os ition to Da rw in 's Fi ve Theories

as in t ran s fo rmational evqlution , but a new beginning is made in

every generation . In deed , evolution is a two-step pheno menon,
the first s tep in each generation being res ponsible for the produc­
tion o f varia tion , which is then so rted in the second step , selec­
tion p roper (see Chapter 6) . Thus , s trictly speakin g , Darwinian
evolution is d iscontinuous because a new start is made in every
genera tion w hen a new set of individuals is produced . That evo­
lu tion nevertheles s a ppears to be to tal l y gradual is becau se it is
population a l and depends on sexual reprod uction a rnong the
members of the popu lation . Such evolu tion is not necess arily
pro g ressive ; it is an opportunistic res ponse to the m o 1nent; hen ce ,
it is unpredictable.

Darwin's Growing Con1 n1itn1ent to Gradualism

The concept of t h e gradual tran sfonn a tion of a population was
not enti rel y new with Darwin . The occu rrence of continuity had
been stressed by son1 e au thors as fa r back as A ristotle, with his
p rinciple of plenitude (Lovej oy 1 9 3 6) . I t was reflected in the con­
cep t of the sea/a natu rae, and even such an arch-es sen tialist as Lin­
naeus stated that the o rders of plants were touching each other
like cou ntries on a map . La n1arck was the first person to apply
the principle of g radualis m to the ori gin of the hierarchy of life ,
bu t there is no eviden ce t h a t Darwin derived h i s g radualistic
thinking fro m Lamarck .
Then how did D a rwin arri ve a t the con cept of gradual evolu­
tion? Referen ces to gradual changes are scat tered through Dar­
win's notebooks from early on ( Kohn 1 9 80) . For ins tance, Dar­
win cons idered the changes of organisms either to be produced
directly b y the environ men t o r to be at least an ans\ver to the
ch anges in the environ ment. Hence, "The chan ges in s pecies
n1 ust be very s low, owing to physical changes s low " (Notebook
C: 1 7) . Gra dualness was also favored by Darwin's concl usion that
ch an ges in habit or behavior n1 ay precede chan ges in structure
(Notebook C: 5 7 , 1 99) . A t that tin1e Da rwin still believed in a prin­
ciple called Yarrel l 's Law (na 1n ed after William Ya rrel l , a natu ral-

. 44 .
 Idcol oJ? ical Oppositio11 to Darw i11 's Fi r,c Theo ries

ist and anin1al breeder), according to which it takes many gener­

ations for the effects of the environment or of use and disuse to
become strongly hereditary. As Darwin stated, "Variety when
long in blood, gets stronger and stronger" (No tebo o k C: 1 3 6). Var­
ious other sources for Darwin's gradualist thinking have been
suggested in the recent literature, such as the writings of J. B.
Sumner (Gruber 1 974: I 25), or Leibniz's principle of plenitude
( Stanley 1 9 8 1 ). But to n1e it scen1s more likely that Darwin ar­
rived at his gradualisn1 owing to two major influences. One was
Lyell's uniforn1itarianis1n, which Darwin extended fron1 geology
to the organic world. The other influence consisted of his own
empirical researches.
At least three observations n1ay have been influential: ( 1) the
slightness of the differences an1ong the 1nockingbird populations
on the three Galapagos islands and the South A1nerican n1ain­
land, as well as a similarly slight difference among many varieties
and species of animals; (2) the barnacle researches, where Darwin
complained incessantly about the extent to which species and va­
rieties were intergrading; (3 ) Darwin's work with races of domes­
tic pigeons, where he convinced himself that even the most ex­
treme races (which, if found in nature, would be unhesitatingly
placed by taxonomists in different genera) were nevertheless the
product of painstaking, long-continued, gradual, artificial selec­
tion. In his Essay of 1 844 Darwin argues in favor of gradual evo­
lution by analogy with what is found in doinesticated anin1als
and plants. And he postulates that "there must have existed inter­
mediate forms between all the species of the same group, not dif­
fering more than recognized varieties differ" (p. 157).
Finally, Darwin had didactic reasons for insisting on the slow
accumulation of rather small steps. He answered the argument of
his opponents that one should be able to "observe" evolutionary
change owing to natural selection by saying: "As natural selection
acts solely by accumulating slight successive favorable variations,
it can produce no great or sudden modifications; it can act only
by very short and slow steps" ( I 859 :4 7 I ) .
Thus, Darwin's rejection of csscntialism and the general en1cr-

• 45 .
 Ideol oJ? ical Opposition to Da rw in 's Fi ve Theories

gence of population thinking strengthened his adherence to grad­

ualism and led him to totally reject saltations. As soon as one
adopts the concept that species evolve as populations and are
transformed, owing to the differential reproductive success of
unique individuals over generations-and this is what Darwin
increasingly believed-one is automatically forced also to believe
that evolution must be gradual. Gradualism and population
thinking probably were originally independent strands in Dar­
win's conceptual framework, but eventually they reinforced each
other powerfully, just as essentialisn1 and saltationisn1 reinforced
one another in the thinking of n1any of Darwin's opponents.
Darwin's totally gradualist theory of evolution-that not only
species but also higher taxa arise through gradual transforn1a­
tion-immediately encountered strong opposition. Even some
of Darwin's closest friends were unhappy about it. T. H. Huxley
wrote to Darwin on the day before the publication of the Or(gin :
" You have loaded yourself with an unnecessary difficulty in
adopting Natu ra non _facit sa ltu m ( Nature makes no jumps] so un­
reservedly" (Huxley 1 900 2:27). In spite of the urgings of Huxley,
Galton, Kolliker, and other contemporaries, Darwin insisted on
the gradualness of evolution, even though he was fully aware of
the controversial nature of this concept. Furthermore, Darwin's
adherence to gradualism became stronger with time; eventually
(after the 1 86 7 critique by F. Jenkin) he minimized even more the
evolutionary role of drastic variations ("sports").
But essentialisn1 and saltationism continued to be widely
adopted. After Darwin's death the concept of gradualism became
even less popular than it had been in Darwin's own time. This
began with William Bateson's I 894 book and reached a climax
with the n1utationist theories of the Mendelians (see Chapter 9).
Both Bateson and de Vries missed no opportunity to make fun of
Darwin's belief in gradual evolution and upheld instead evolution
by macromutations ( Mayr and Provine 1 9 80). A n1ild popularity
of saltationist theories continued right through the evolutionary
synthesis (Goldschmidt 1 940; Willis 1 940; Schindewolf 1 950).
The naturalists were the n1ain supporters of gradual evolution,
 Jdeolo<�ical Opp osition to Da rw in 's Fi t,e Th eo ries

which they en coun tered everywhere in the form of geographic

varia tion . Eventually, geneticists arri ved at the same con clusion
through the dis covery of ever sli gh ter mutation s .
Defining gradualism as populational evolution - and this is
what Da rwin b asically had in mind - permits us to say that in
spite of all the op position to hi m , Darwin ulti matel y prevailed
even with his fou rth evol utionary theory. The only exceptions to
gradualisn1 that are clearl y establis hed are cases of stabil ized hy­
brids that can reproduce wi thou t crossing .
Nothing is said i n the theory o f gradu alis m about the precise
rate at which the ch ange may occur. Da rwin wa s aware of the fact
that evolu tion could so meti mes progress rapidly, but, as An drew
Huxley ( 1 9 8 1 ) has poin ted out, evolu tion could also con tain pe­
riods of co m plete stas is " during which these same species re­
mained without u ndergoing any change ." I n a well-known dia­
gram in the Origin , D a rwin lets one species (F) con tinue
unchan ged th rough 1 4 , 000 genera tions or even through a whole
series of geologica l strata . Understanding the independen ce of
gradu alness a n d evolu tionary rate is im portant for evaluating the
theory of punctuated equilibria put fo rth in 1 972 by Stephen J.
Gould and N iles Eldredge (see Cha pter 1 0) .

• 47 .
 C H ·A P T E R F I V E
-------------•-------------

The Struggle against Ph y sicists

and Philosophers

E SSEN TIA LI S M W AS NOT the only ideology Da rwin had to

overcom e . A concept of science had developed sin ce the
seven teen th cen tury that was co mpletely do n1inated by physics
and mathem atics . The philoso ph ers fro 1n Bacon aiid Descartes to
Locke and Kant en tirel y ag reed with the phys ical scien tists fro m
Galileo a n d New ton t o Lavoi sier a n d Laplace that the ideal o f sci­
ence should be to establish n1 athen1atically fo rmed theo ries that
were based on universal laws . The possibility of proof and of ex­
act p rediction were the tes ts fo r the goodness of a scientific ex pla­
nation . Newton's ph ysics was the s hining exam ple of good sci­
ence . E very B ritish scientist and philosopher, at the time when
Darwin was developing his ideas on evolution , agreed in this
concept of science, and there are indi ca tions that Dar win did his
best, particularl y after reading and rereading John Hers chel 's Dis­
course , to Ii ve up to his ideal ( Ruse 1 979) .
In spite o f all these efforts , Darwin's empirical res earches led to
res ults that were in conflict with most of the basic assumptions of
phys icali s m . The ph ysica lists were essentialis ts , a philosophy that
Darwin to tally rej ected . Instead, Da rwin developed population
thinkin g , a n1 ode of thinking utterl y alien to the physicalists . The
physicists at that time were strict determinists ; prediction was not
only possible but was the ve ry tes t of the validity of theo ries .
Evolutiona ry p rocesses , by con trast, involved a considerable
cha n ce elen1en t : they were p robabilistic, and hence they did not
pennit absolute p rediction . All evolutionary pheno1nena could
 The Strug_�le aga inst Ph ysicists and Ph ilosophers

be explained on ly by inferrin g past historical even ts , a consi dera­

tion absent (at that period) in the physical sciences . The probabi­
listic nature of the evolutionary process was so alien to the think­
ing of the ph ysicists that H erschel referred to natural selection as
the theory of the " higgledy-piggledy " (F. D a rwin 1 8 8 8 , 2 : 240) .
Darwin's findings co 1n pletely undern1 ined the physicalist con­
cept of la ws . The order and harmony of the crea ted universe
made the ph ysical scientists sea rch for laws , fo r wise reg ulations
in the running of the universe ins talled by the C reator. To serve
his Creator b est a physicist studied His laws and their workin g .
In this tradition Darwin refers in the Origin o.f Species n o fewer
than 1 06 times in 490 pages to laws controlling certain biological
p rocesses . B u t Darwin's " laws " were not the laws of the deists
but were either s i m ple facts or regular processes . No longer re­
lying on univers al laws , D arwin had no problem in accepting sta­
tistical generalizations . I t was a co mplete rej ection of Cartesian­
Newtonian determinis m .
The role D arwin assigned to chance has never been properly
analyzed . The deterministic spirit of science at his time was in
complete conflict with Darwin's findings, which showed how
strong a role in evolu tion was played by chance. In the case of
variations , he dis tinguished between those being accidental as far
as their " purpo�e" or selective value is concerned , and others that
are " accidental as to their cause o f origin " (F. Darwin 1 8 8 8 l : 3 1 4) .
A similar train o f thought is expressed in the Variation of Animals
and Plan ts ( 1 8 6 8 2 :4 3 1 ) . I t is evident that Darwin accepted the
strict working of what he called natural laws at the physiological
level but was aware o f chance (stochastic) processes at the orga­
nis mic level .
The realization that neither essentialism nor determinism nor
any other as pect o f physicalism was a valid ideology was of the
utmost i m portance for the further development of Darwin's
though t . Darwin could have never adopted natural selection as a
maj o r theory, even a fter he had arrived at the principle on a
largely empirical basis, if he had not rej ected essen tialism and

• 49 .
 The Struggle aga inst Ph ysicists and Ph ilosophers

physicalism. But there was· one other impeding ideology Darwin

had to refute in order to be able to adopt natural selection, and
that was the finalistic or teleological worldview.

Final Causes
Again and again the statement has been made that "Darwin \Vas
no philosopher," even by an otherwise so perceptive author as
G. G. Simpson ( 1 964: 5 0). In fact, Darwin was keenly interested
in philosophy and, as we have seen, atten1pted to follow in his
own writings the best advice of the philosophers of science of his
day. Admittedly, he never published an essay or volun1c explicitly
devoted to an exposition of his philosophical ideas, but in h is
scientific works he systematically demolished one after the other
of the basic philosophical concepts of his time and replaced them
with revolutionary new concepts.
In retrospect it is rather surprising to what an extent the con­
temporary philosophers who were involved with scientific ideas
ignored Darwin or at least failed either to incorporate his con­
cepts into their own systems or at least to try seriously to refute
him adequately. This is as true for the British philosophers Her­
schel, Whewell, Mill, and Jevons, as well as for the various Ger­
man philosophers and philosophical biologists, the followers of
Kant, the later Naturphilosophen , the mechanoteleologists (Lenoir
1 982), and the mechanists (DuBois-Reymond, Helmholtz,
Sachs, Ludwig, Loeb). To a large extent these philosophers failed
to recognize the importance of Darwin's ideas because of their
co1nmitn1ent to the philosophies of essentialism and finalisn1.
From the days of the earliest philosophers it was widely be­
lieved that the world must have a purpose because , as Aristotle
had said, " Nature does nothing in vain," and neither, a Christian
\vould say, does God. Any change in this \vorld, they would say,
is due to final causes that n1ove the particular object or phcnon1-
enon toward an ultin1ate goal. The developn1ent of an organisn1
fron1 the fertilized egg to the adult stage \Vas frequently cited�
fron1 Aristotle on, as an illustration of this striving to\vard a goal .

. 50 •
 Th e Strug�lc aga inst Ph ysicists and Ph ilosop h ers

I t wa s a l n1 o s t universally believed that everything in na ture , par­

ticu larly all d i rectional proces ses , 1noved in an anal ogous manner
towa rd a p redetcnn ined en d . Those who adhered to this view
have been designated teleologists , or " fi nalists . "
T h e thinkers o f t h e S cientific Revolution i n the six teenth and
seven teen th centu ries \Vere fascinated by n1 otion . They worked
o u t the laws of fal lin g bod ies and the n1 otion of the planets
around the sun ; fo r then1 , the wo rld was a wo rld of 1notion con­
trolled b y eternal l a ws . God had instituted thes e laws at the time
of c rea tio n , but fro m that point on laws kept the world moving .
God was the final cause o f everythin g , but H e ru led the wo rld
through H is laws and not b y continuous interven tion . Rene Des­
cartes was one of the chief s pokes n1en fo r this s t rictl y physicalist
or m echan istic wo rl d v iew, which was pri m a rily that of the phys­
icists . It was , howe ve r, more or les s adop ted even by the natu ral­
ist B u ffo n a n d carried to the n1 ost extre n1 e con sequences by
Ba ron H olbach . Even those who adopted this concep t of a mech­
anized world had certain mis givings about applying i t to the liv­
ing wo rld . B u ffon , fo r ins tan ce , was fu lly aware of the conflict
between the mechanized wo rld p icture and many pheno mena en­
countered in the study of organ is m s . Yet any alterna tive view was
unaccep table to h i m .
Those w h o were dissatisfied with a st rictly mechanistic world ,
entirely ru n b y laws , developed a differen t explana tory frame­
wo rk . They credited God with a much la rger role in designing
the world dow n to the last detail and in effectin g the changes that
had taken place sin ce creatio n . I t was distas teful to them to re­
m ove God fro m the running of His wo rld and to replace Him b y
t h e efficien t cau ses o f His l a ws . Not o n l y that, but they also
found it in con cei vable tha t the observed harmony of nature and
all the m u tu a l adap tations o f organisms to one another could be
due si m pl y to efficien t causes . Thei r answer wa s to stress the elab­
orateness o f the ori ginal design of the worl d to a fa r greater de­
g ree than had been done by the mecha nists . No n1 at ter where yo u
loo ked in nature, they clai med , you would find evi dence for the
in finite wisdom of the Creator. Anyone who wou ld study His

. 51 •
 Th e Struggle against Ph y sicists and Ph ilosophers

work (nature) was as legitimate a theologian as he who would

study His word (the Bible) . Beginning with John Ray ( 1 69 1 ) and
William Derham ( 1 7 1 3 ) , the study of na ture became physico­
theology or natural theology. I t became the study of design .
Two further developments strengthened the belief in final
causes . One was the in creasingly strong belief that God had cre­
ated the world for the sake o f man . This was foreshadowed by
Aristotle's statement (Politics 1 , 8, 1 2 56a , b) , " Now if nature
makes nothing incomplete and nothing in vain , the inference
must be that she has made all animals for the sake of man . " It was
made legitimate by corres ponding statements in Genesis . The
other reinforcement of the belief in final causes came from mani­
fold observations indicating ongoing changes in the wo rld. This
led to a new concept of creation . Creation was no longer seen as
so mething that had happened instan taneously (or in six days) , but
as a gradual and slow process, directed by final causes , cul minat­
ing in the production o f man . Consistent with this modified con­
cept of creation, G . W. Leibniz and J . G . Herder temporalized the
sea /a naturae, which more and more was considered a scale of per­
fection . One of the foremos t obj ectives of the writings of the
physicotheologians was to demonstrate how perfectly everything
in the wo rld was designed . Since God could not have created any­
thing that was not perfect , the world was considered the " best of
all possible worlds . " This was a do minant theme of that vas t lit­
erature from Ray and Derham to William Paley and the Bridge­
water Treatises . It dominated even Darwin's early thinking and
certainly that of most of his contemporaries . Until evolu tion was
accepted , there was no conce� vable alternative to chance but "ne­
cessity, " that is , God 's design .
Much of the literature of natural theolo gy is quite admirable .
R . Boyle ( 1 6 8 8) , for ins tance, understood perfectly \vell that the
explanation of the mechanical workings of a stru ctu re is an en­
tirely independen t endeavor from the explanation of the reason
why the organ exists and what its role in the life of the organism
is . Thus , he made quite clearly a dis tinction bet\veen proxin1ate
or i1n n1ediate causations and ulti n1ate causations . For instance ,

. 52 •
 The Struggle aga inst Ph ysicists and Ph ilosoph ers

the proximate cause of sexual dimorphism in the plumage of

birds is horn1onal difference; the ultimate causation is sexual se­
lection. Proximate causations could be explained mechanistically,
by physical laws, but one could not explain ultimate causations
without postulating a final goal or purpose (Lennox 1 98 3 ) .
Though the beginnings of natural theology go back to the
Greeks and even the Egyptians, its period of true do1ninance , at
least in England, lasted from the last quarter of the seventeenth
century to I 8 5 9 . It made little difference whether an author be­
lieved that everything in the world was governed by laws or was
specifically regulated by God, because in either case God was
either directly or indirectly responsible. He was the final cause of
everything.
A belief in cosmic teleology fit well into the thinking of the
seventeenth and eighteenth centuries . This was a period of in­
creasing optimism, of emancipation from social and legal bur­
dens, of conviction that better times were coming, possibly a mil­
lennium. Progress was preached not only by utopians and
reformers but became the theme of philosophies, particularly of
the historical-idealistic schools from Herder and Schelling to He­
gel and Marx (Toulmin 1 98 2) . Nowhere else did teleology have
as great an influence as in Germany. Almost all German philoso­
phers, from Leibniz, Herder, and Kant to modern times, were
teleologists to a greater or lesser extent. Kant, whose thought
dominated German philosophy throughout the nineteenth cen­
tury, was a teleologist (Low 1 9 80) . As far as inanimate nature was
concerned, he was a strict mechanist, but he considered all phe­
nomena of living nature to be the product of teleological forces.
Under Kant's influence, German biology in the first half of the
nineteenth century was permeated with teleological thinking
(Lenoir 1 9 8 2) . K . E. von Baer's comprehensive critique of Dar­
win was largely based on teleology, and so was that of the philos­
opher Eduard von Hartmann ( 1 8 76) . And the post-Darwinian te­
leological theory of orthogenesis had nowhere as 1nany followers
as in Germany.
Teleological thinking was strongly reinforced by the studies of

• 53 •
 Th e Struggle aga inst Ph ys icists and Ph ilosophers

the geologists and particularly b y the dis covery of succes sions o f

fos s il faunas cul m inating i n strata containing mammals a n d even­
tually man ( B owler 1 9 76) . It fit well with Lamarck 's theory o f
gradual evolutionary chan ge, this being the first genuine theory
of evolution ( 1 809) . Not all progressionis m in geology led to the
acceptance of evolu tion ; in fact , the maj o rity of paleontologists
fro m C u v ier to A g assiz thought, rather, in terms of catas trop hes
and s u bsequent more progressive new creations . Fewer and fewe r
autho rs continued to insist on the cons tancy of the wo rld ; n1ost
of them saw continuous change and indeed a trend toward per­
fection . This can be perceived in the writings of al most all au thors
between I 809 and I 8 59, even though it was expressed in various
ways b y authors like M eckel , Chambers , Owen , Bronn , von
B aer, and A gassiz .
The general opti111 isn1 o f the ei gh teen th cen tu ry received a se­
vere j olt th rou gh the disastrous Lis bon earthq uake of Nove 1n bcr
1 , 1 7 5 5 . It induced Voltaire to satirize the Pang lossian thin king of
A lexander Po pe and Leibniz in his Candide . David Hun1c also rid­
iculed claims of a hann ony of nature : " Ins pect a little 1n ore nar­
row l y these l i ving existences , the only beings wo rth reg ardin g .
How hos tile a n d destructive ; how insu fficien t a l l of then1 fo r their
own happiness ! How conten1 p tible or odious to the spectato r !
The whole p resen ts nothin g but the idea of a blind nature . " Kant
likewise refuted the claims of natu ral theology. The unhappy
conseq uen ces o f the French Revolu tion con tribu ted to the
sprea din g of a deep pessimisn1 . It is reflected in the thinking o f
Malthus a n d other demographers . No longer was t h e grow th o f
hu n1 a n popula tions seen a s on e of t h e benefits bes towed on n1 an
by God . Ra ther, it was clai med that ow ing to li111its in1posed by
the environ n1ent , such grow th would inev itably lead to poverty,
disaster, and death .
The 1nore the s tudies of the naturalists progressed, the 1nore
often phen o n1ena we re fo und that cont radicted the excellen ce of
desig n . Not every organisn1 could have been exclusivel y designed
fo r its role in n a tu re : how would this account fo r the exis tence of
a l i 1nited nun1ber o f well-defined ty pes , such as n1 an1 n1 als , bird s ,

• 54 .
 Th e Strug� le against Ph y sicists and Ph ilosophers

sna kes , beetles , and s o on ? Ra ther, it wa s said, at the beginning

rel atively fe w archetypes were crea ted and the laws of nature g ave
rise to the su bsequent diversity, everyth in g , howeve r, had been
contained in the plan of crea tion . Th us, indirectly, even in this
thin k in g , diversity and adaptat ion were due to des ign ( B owler
1 9 77) .
B u t this revision o f the design argu men t cou ld not silence crit­
i c i s n1 . One a s ked : What is s o wonderful about a parasite tha t tor­
tures its v icti n1s and leads to their even tual death? Even wo rse,
how could design be p erfect i f it leads to such wides pread extinc­
tion , a s docu m en te d b y the fo ssil record ? I f the h a nnony of the
living wo rld, as descri bed b y n a tu ral theologians, is reflected by
the m u tu a l ada ptation of organisms to one another and to thei r
env i ro nm en t , and i f these adapta tions must be adj usted continu­
ous} y to cope with the ch anges o f the earth and with the res truc­
turi n g o f the fa unas ow in g to ex tinction , what final causes coul d
there be to govern all these ad h o c chan ges ? I f the envi ronmen t
chan ges , the organi s m has to readj u s t to i t . But there is no nec­
essary direction , n o thought of necess ary progres s , and no rea ch­
ing of any final g oa l s . A fter evolutionary thinking had begun to
s p read, Matthias S chleiden ( I 8 42 : 6 1 ) insisted that although one
can observe sim ple as wel l as com plicated organi s m � , " it would
be a totally misleading language i f we would use for them the
words i m perfect and perfect, or lower an d higher. "
Natura l theolo g y, with i t s em phasis o n design, had been vir­
tua l l y a ban d oned on the Con tinen t b y abou t I 800 . But it contin­
ued to be s trong in England , an d all of Darwin's teachers and
peers , p a rticularly Sedgwick, Hensl ow, and Lyell , were con­
firmed natura l theo logian s . This was Da rwin's conceptual frame­
wo rk when he began to think about adaptation and the origin o f
s pecies .

From Natural Theolo g y to Natural Selection

There are m any in dica tions that when D a rwin returned from the
BeaRle voyage he shared the beliefs of the na tural theologian s . He

· 55 ·
 The Struggle aga inst Physicists and Ph ilosophers

had wholly abandoned them, twenty-three years later, when he

published the Or(gin . There is, however, not yet complete con­
sensus in the Darwin literature under what influences and in what
stages Darwin revised his interpretations. It was a peculiar pe­
riod, since the British philosophers of science-Herschel,
Whewell, and Mill-emphasized a rigorous scientific methodol­
ogy and yet all firmly believed in final causes. They believed in
laws, but God's guiding hand was needed because unguided laws
would lead to random disorder (Ruse 19756; 1979).
To what did the young Darwin attribute adaptation ? Prior to
1 83 8 his ideas on this point were rather vague. He seen1s to have
attributed adaptation to certain laws, particularly the influence of
the environment on the generative system. He still thought in
terms of the design of the world. In his Transmutation Notebooks
of this period, the most clearly teleological staten1ent refers to
dispersal: "When I show that islands would have no plants were it
not for seeds being floated about , -1 must state that the mecha­
nism by which seeds arc adapted for long transportation, seems
to imply knowledge of whole world-if so doubtless part of sys­
tem of great harmony " (D: 74). Darwin's pre- I 8 3 8 interpretation
of evolutionary change depended on God's planning and was thus
clearly a finalistic interpretation. For the Dar\vin of the Transmu­
tation Notebooks (before September 1 83 8), the seen1ing path of
progression toward perfection was simply the result of certain
laws that n1ade such a development possible. All organic change,
he thought, was an adaptive response to changes, however slight,
in external conditions. These environmental influences induced
the generative system to produce appropriate responses. This im­
plied that God was directly involved in adaptation because only
God could have 1nade the generative system in such a way that
changes in the environn1ent would induce it to con1e up \Vith an
adequate response.
Yet, as l)arvvin's studies proceeded he discovered one pheno111-
cnon after the other that cast doubt on the perfection of adapta­
tions (Ospovat 198 1 ). First, he discovered all sorts of evidence for
,
descent ( called "propagation , or progression in Dar\vin's earlier
 The Strug(!le a(_(? a inst Ph ysicists and Ph ilosophers

notes) , which served as a definite constraint on the absoluteness

of adaptation. Then came the consideration of rudimentary or
vestigial organs, which also contradicted perfect adaptation, as
did the widespread occurrence of extinction. Those natural theo­
logians, and there were others beside Darwin, who saw such in­
consistencies and seen1ing incon1patibilities with the concept of a
total harn1ony of nature ascribed the deviations fron1 perfect ad­
aptation to a conflict between various laws instituted by the Cre­
ator. Organisms, said these authors, were only as perfect as is
possible within the lin1its set by the necessity of conforn1ing to
these laws. There are, for instance, different laws required to ex­
plain the facts of structure, distribution, and succession.
Somehow such a direct reliance on eternal God-given laws for
the explanation of natural phenomena must have been unsatisfac­
tory to Darwin and in conflict with some part of his major phil­
osophical framework. This must be the reason why he aban­
doned this type of thinking so speedily after he read Malthus and
formulated his theory of natural selection on September 1 8 , 1 83 8.
Natural selection gave him a purely mechanistic explanation for
adaptation and for evolutionary progression. As Darwin stated in
his A utobiography ( 1 9 5 8 : 8 7) : "The old argument of design in na­
ture, as given by Paley, which formerly seemed to me so conclu­
sive, fails, now that the law of natural selection has been discov­
ered. We can no longer argue that, for instance, the beautiful
hinge of bivalve shell must have been made by an intelligent
being, like the hinge of a door by n1an. There seems to be no
more design in the variability of organic beings and in the action
of natural selection, than in the course which the wind blows. "
After 1 8 3 8 Darwin at first remained enough of a natural theo­
logian to believe that natural selection could give him perfect ad­
aptation. But, he abandoned this belief by the 1 8 5 0s, and the Ori­
gin is remarkably free of any teleological language (Ospovat
1 9 8 1 ). To be sure, the word "progress" is used ten times in this
volume but almost always as a term to describe a passing of time.
Only in connection with the replacement of fossil faunas of
which each one seems to be "higher" than the one it has replaced

· 57 ·
 The Struggle aga inst Ph ysicists and Ph ilosophers

does Darwin speak of a protess of improvement; but he adds, "I

can see no way of testing this sort of progress" ( I 8 59 : 3 3 7). How­
ever, Darwin points out that there are differences in competitive
ability even among living faunas. British fauna} elements intro­
duced to New Zealand are highly successful, while he doubts the
reverse would be true. "Under this point of view," says Darwin,
"the productions of Great Britain may be said to be higher than
those of New Zealand." Yet this is not a teleological argument.
The greater competitive ability of the faunal elements of Great
Britain was not due to any built-in drive or final cause but simply
due to the fact that the British fauna had passed through a n1ore
severe struggle for existence.
Nevertheless, the concepts of "perfect" and "perfection" con­
tinued to be popular with Darwin. In the Or((!in he used the word
"perfect" 7 7 times, "perfec ted" 1 9 tin1 es, and "pc rfe c tion" 2 7
tin1es. What is remarkable, however, in these uses is how care­
fully Darwin makes a distinction between the product of selec­
tion and the process of perfecting. We look to his explanations in
vain for a drive or tendency toward perfection. Invariably Dar­
win en1phasizes that selection carries the evolutionary line to
ever-greater perfection. This is particularly well-stated in the sec­
tion of Chapter 6 with the heading "Organs of Extreme Perfec­
tion and Complication" (p. 1 8 6 ) which, among others, contains
Darwin's well-known discussion of the evolution of eyes through
natural selection. Since natural selection is not a finalistic process,
Darwin now sees quite clearly that "natural selection will not
necessarily produce absolute perfection; nor as far as we can judge
fro1n our limited faculties, can absolute perfection be everywhere
found" (p. 206 ) . Complete perfection, of course, is not needed,
because "natural selection tends only to make each organic being
as perfect as, or slightly n1ore perfect than, the other inhabitants
of the san1e country with which it has to struggle for existence.
And \Ve see that this is the degree of perfection attained under
,
nature . (p. 20 1 ) . There is not even a trace of a suggestion of any
final cause, because perfection is sin1ply the product of the a pos­
terio r process of natural selection. With the \vorld and its biota
constantly changing, perfect creation in the beginning \Vould
 Th e Stru(_(!gf e aJ?a inst Ph ysicists and Ph ilosophers

have been futile. "Almost every part of every organic being is so

beautifully related to its complex condition of life that it scen1s as
improbable that any part should have been suddenly produced
perfect, as that a complex n1achine should have been invented by
man in a perfect state" (Origin , 6th ed. , pp. 58 -59).
Darwin's subsequent correspondence with the An1crican bota­
nist Asa Gray pern1its us to analyze his thought even a little fur­
ther. Gray, even though a rather strict creationist, accepted the
importance and guiding capacity of natural selection. However,
"Natural selection is not the wind which propels the vessel, but
the rudder which, by friction, now on this side and now on that
shapes the course" ( Moore 1 979:3 1 6). Variation-the wind in
Gray's metaphor-was guided for Asa Gray by a divine hand.
This possibility was emphatically rejected by Darwin and in­
duced him to state his ideas, rather evidently as a direct answer to
Gray, in Th e Va riation of A n imals and Plan ts ( 1 86 8 I l :43 2). Gray,
however, failed to understand Darwin's argument and went even
so far as to praise " Darwin's great service to natural science in
bringing back to it teleology" (Gray I 8 76 : 23 7).
In his later years, particularly in letters to his numerous corre­
spondents, Darwin was sometimes rather careless in his lan­
guage. For instance, he referred to "the extreme difficulty or
rather impossibility of conceiv ing this immense and wonderful
universe, including man with his capacity of looking far back­
wards and far into futurity, as the result of blind chance or neces­
sity." How could he have said this when the theory of natural
selection had given him exactly the means to escape from the al­
ternatives chance or necessity ? On another occasion he said, " The
mind refuses to look at this universe, being what it is, without
having been designed." It is not surprising therefore that Darwin
was misclassified by a number of authors who did not understand
the working of natural selection. Kolliker, for instance, accused
Darwin of being "in the fullest sense of the word a teleologist. "
And even T. H. Huxley when defending l)arwin was driven to
distinguish between "the teleology of Paley and the teleology of
evolution" (Moore 1 979: 264).
Darwin was not totally alone in his rejection of finalisn1. Ernst

. 59 .
 The Struggle against Physicists and Ph ilosophers

Haeckel declared emphatic'illy that " the causes of all phenomena

of nature . . . are purely mechanically acting causes , never final,
the goal-directed causes " ( 1 8 66 2 : 1 5 0) . The most articulate
among D arwin's supporters was August Weismann, who took up
the battle for natu ral selection again and again and refuted the
theories of D arwin's opponents (see Chapter 8) . The voices of
Haeckel, Weis mann , F. Miiller, and Darwin's naturalist friends
were, however, merely cries in the wilderness, for the opposition
to the mechanistic process of natural selection was almos t univer­
sal . But none of his op ponents truly unders tood natural selection ,
and this misunderstanding was to a large exten t due to a long­
standing ideological co mmitment to finalism . The opposition to
natural selection continued up to the evolutionary synthesis and
with it an open or uns poken support of finalis m . For exam ple,
the geneticist T. H. Morgan , who showed his lack of understand­
ing of natural selection even in his last book on evolution in 1 9 3 2 ,
clain1ed in 1 9 1 o that finalism had entered biology through natu ral
selection because " b y picking out the new va riation . . . purpose
enters in as a factor, for selection had an end in view, " completely
ignoring the rando n1 ness of variation and the statistical nature of
the selection process .

Fin ali s m as a n Alternative to Natural Selection

N umerous attempts were made in the years after 1 8 5 9 to replace
Darwin's theory of natural selection with a superior way of
achieving adaptation. The best-known of these theories are usu­
ally classified under the headings neo-Lamarckisn1 (inheritance of
acquired characters) , orthogenesis (an intrinsic perfecting prin­
ciple) , and saltation . They all incorporated some finalistic com­
ponents to a lesser or greater extent. It is not easy to report on
these theories for a nu mber of reasons . Not only are the descri p­
tions o f the postulated n1echanis m by which the changes are
achieved usually quite va gue, but the same author may support
first one and then the other of these theories , or a n1ixture of then1
(K ellogg 1 907; Bo\vler 1 98 3 , 1 9 8 8 ) . Even after the Paleyan con­
cept of an ad hoc design of every-even the slightes t-adaptation

. 60 .
 Th e Struggle aga inst Ph ysicists and Ph ilosophers

had lost all credibility, there remained a concept of a universal

design of organic progression, an evolutionary reinterpretation
of the temporalized scale of nature (Bowler 1 977) . Such a concept
seemed, at first, to have a sound observational foundation. Con­
sidering that variation is random, as Darwin postulated, and con­
sidering that the nun1ber of environn1ental constellations is quite
unlimited, one would expect a totally chaotic network of evolu­
tionary phenomena . What one actually finds is the existence of a
limited number of well-defined lineages and the possibility of ar­
ranging organisn1s into progressive series. This was described
not only b y paleontologists but also by students of living orga­
nisms, be they butterflies (Theodor Eimer) or birds (Charles 0 .
Whitman) . Variation evidently was not random but followed
well-defined pathways of change. Such evolutionary trends were
ascribed to an intrinsic, direction-giving force, called orthogene­
sis. It was described as a perfecting principle or (in German) Ver­
vollkom,nnungstrieb.
The intrinsic nature of this force seemed to be confirmed by
the fact that it was possible to establish rectilinear series not only
for characters that might have been advanced by natural selection,
such as increasing precision of mimicry patterns or phyletic in­
creases in body size, but also for nonutilitarian or seemingly del­
eterious characters. This was an argument made particularly en1-
phatically b y Eimer (Bowler 1 979) . Most proposals of ortho­
genesis were made in strict opposition to and as alternatives to
natural selection.
However, there was a group of Christian Darwinians for
whom natural selection was "evidence of a directing agency and
of a presiding mind" (Moore 1 979) . They either thought that var­
iation as such was directive, supplying just the right material to
selection, or they considered the selecting process as purposive .
Clearly for them natural selection was a teleological process .
Even such an ultra-mechanist as Julius Sachs ( 1 894) adopted
Carl Naegeli's perfecting principle as the agent of all major evo­
lutionary developments, with natural selection n1erely being able
to improve fine-grained adaptation . Kollikcr ( 1 864) was another
adherent to an autogenetic theory ascribing all evolutionary pro-

. 61 •
 Th e Struggle aga inst Ph ys icists and Ph ilosophers

gress to "intrinsic causes," · lnd like Naegeli he stimulated Weis­

mann to a reply.

Evolutionary Progress without Final Causes

For some Darwinians the concept of evolutionary progress seen1s
to have raised some embarrassing questions. How can a strictly
opportunistic competitive struggle lead to progress? l)arwin
himself occasionally seems to have had such doubts, and they are
reflected in his comment, "Never use the words higher or lower,"
written on the margin of his copy of the Vestig es . Others who also
questioned progress pointed to the continued existence of the ar­
chaebacteria and other prokaryotes, to the great flourishing of the
protists and lower fungi up to the present, to the parasites, and to
the inhabitants of caves. None of these can be called progressive,
in the sense of "higher," and yet they continue to exist and flour­
ish. So, it was said, evolution is sin1ply a process of specialization.
And yet, who can deny that overall there is an advance from the
prokaryotes that dominated the living world more than three bil­
lion years ago to the eukaryotes, with their well-organized nu­
cleus and chromosomes as well as cytoplasmic organelles; fron1
the single-celled eukaryotes to plants and anin1als with a strict
division of labor an1ong their highly specialized organ systems;
and, within the animals, from ectotherms that are at the rnercy of
clin1ate to the warn1-blooded endotherms; and, within the en­
dothern1s, from types with a small brain and low social organi­
zation to those with a very large central nervous system, highly
developed parental care, and the capacity to transn1it inforn1ation
frorn generation to generation?
Atten1pts to define progress have been n1any. For Lamarck, for
instance, and for many nineteenth-century authors man was
clearly the rnost perfect organisrn, and all forn1s of life \Vere ar­
ranged in a single column on the basis of their assumed progress
toward rnanhood. No\v we know that diversification is the n1ost
characteristic attribute of evolution and that life not onlv vcrv ,I .,

early split \vithin the prokaryotes into eubacteria and archaebac-

teria, but the eukaryotes after their origin quickly gave rise to the

. 62 •
 Th e Strug�le against Ph ysicists a 11d Ph ilosophers

protists, and to the kingdoms of fungi, plants, and animals. Lit­

erally thousands of distinct phyletic lines developed within each
of these kingdo1ns, n1ost of then1 not in the slightest tending to­
ward the characteristics of 1nan. Neither can the don1inancc of a
group on earth be considered the criterion of progress. On that
basis the vascular plants would have to be considered n1ore don1-
inant than man and even the insects. And n1an's ancestors until
less than ten thousand years ago were anything but don1inant on
earth.
Structural con1plexity is son1etin1es mentioned as a sign of pro­
gress, but trilobites and placoderms would seen1 to have been
more complex in structure and perhaps more specialized than
modern man. Huxley ( 1942) considered en1ancipation from the
environment an in1portant index of progressiveness, and in that
criterion man certainly ranks higher than any other organism.
However, is independence from the environment truly an index
of progressiveness?
W hen discussing evolutionary progress one seems to be quite
unable, since one is a member of the human species, to get away
from criteria that would give man supremacy. However, there are
two criteria of progressiveness tha._t would seem to have a consid­
erable amount of objective validity. One of these is parental care
(promoted by internal fertilization) , which provides the potential
for transferring infonnation nongenetically from one generation
to the next. And the possession of such information is of course
of considerable value in the struggle for existence. This informa­
tion transfer generates at the same time a selection pressure in
favor of an improved storage system for such remembered infor­
mation, that is, an enlarged central nervous system. And, of
course, the combination of postnatal care and an enlarged central
nervous system is the basis of culture, which together with
speech, sets humans quite aside fro1n all other living organisn1s.
However, even if we would designate the acquisition of these ca­
pacities as evidence for evolutionary progress, it would not
strengthen the case for final causes, since these developn1ents
were clearly achieved through natural selection.
Whether one is looking at the highest 111a n1111als and birds, the

. 63 .
 The Stru(_� le against Physicists and Ph ilosoph ers

social insects , the orchids , or giant trees , it has seemed inconceiv­

able to some students of evolution that the slow struggle fo r ex­
isten ce among individuals of a species could account for the enor­
mous evolutionary progress observed in so many phyletic lines .
To see all evolution simply as the result of co mpetition among
the individuals in a population is indeed simplistic, because su­
perim p osed on this individual selection is a process traditionally
referred to as species selection , although perhaps a better term
would be species replacement or species succession . An individ­
ual o rganism competes not only with members of its own species
but struggles for existence also against members of other species .
And this process is probably the greatest source of evolutionary
progress . Each newly formed species , if it is evolutiona rily suc­
cessful, must represen t, in some way, evolutiona·ry progress .
Darwin explained this as follows : " B ut in one particular sense the
more recent forms must , on my theory, be higher than the more
ancient; for each new species is formed [ that is , has become suc­
cessful ] by having had son1e advantage in the struggle for life over
other and preceding forn1s " ( 1 8 5 9 : 3 3 7) . When the com petition
among individuals of different species leads to or at least contrib­
utes to the extinction of one of the co mpeting species , it is a case
of species replacement . That com petition an1ong species could
lead to the extinction of one of the competitors was of course
already known to Lyell and other pre-evolutionary authors .
A new species will be su cces sful in the struggle fo r existence
over a previously existing species onl y if it has made son1e, even
the sn1 alles t, evolutionary invention . This n1 ight be an i n1 prove-
111ent in its diges tive physiology or its nervous syste 1n or its life­
style or any other of the countless ways by \vhich the so-called
"higher" organis m s differ from the lower ones . Thu s , the Dar­
\vinian 1nechanis n1s of variation and selection , of speciation and
extinction , are fully capable of explaining all 1nacroevolutionary
developn1ents , whether specialization s, improven1en ts, or other
innovation s . And none of this requires any finalistic agen t .
Close study of evolutionary prog ress shows that its character­
istics arc not co 111pa tible with \vhat one \Votild expect fron1 a pro-

. 64 .
 Tlze Strug(tle a'"(t a inst Ph ys icists and Ph ilosop h ers

cess g uided b y final causes . P rogressive chan ges in the history o f

life a re neither p redictable n o r goal-di rected . The ob served ad­
vances a re ha phaza rd and hi ghly divers e . I t is al ways uncertain
w hether newly acquired adaptations are of perm anent va lue .
Who at the beginn ing o f the Cret aceou s would have predicted the
total extinction by the end of the period of that flourishin g ta x on
the dinosaurs ? E pisodes of s tasis al ternate with epis odes of pre­
cipitous evo l u tionary ch ange. Evolu tion ary trends arc ra rel y rec­
til inea r for any len gth o f t i n1 e , and when such rectilinea ri t y oc­
curs it can u s u a l l y be shown to be due to buil t-i n co nstrain ts .
A l l the evo lutionary phenon1 ena and as pects of evolutionary
p rogress that were considered as irrefutable p roof of teleology b y
ea rl ie r genera tions can now b e shown t o b e en tirely con sistent
with natural selection . Phen o n1ena that are due to a chain of his­
tori ca l even ts cannot be a scribed to sin1 ple laws and can therefore
not be proven in the s a n1e way a s can phenomena stu died in the
physical s cien ces . H owever, they can be sh own to be consis ten t
with the findings of genetics and with the theory of natural selec­
tion in its m odern sophisticated fo nn . No one has refu ted the
finalistic thesis of e volution more convin cingly than Sin1 pson
( 1 949 ; 1 9 74) . He pointed out that each evolu tionary line goes its
own way, and evo l u tion ary p rogres s can be defined only in tern1 s
o f th a t p a rticular lineage. Nothing seemed more progressive in
the geologica l p a s t than the a m monites and the dinosaurs , and
yet both taxa beca m e extin ct . On the other hand m any evolu tion­
a ry lines have dis p l ayed no eviden ce of progress in hundreds or
thousands of mil lions of years , and yet they have s urvived to the
p resent day, as the archaebacteria and other p rokaryotes . Progress
thus is not a t all a univers a l as pect of evolution , as it o ught to be
if evolution were generated by final causes .

The Decline, if not Demise, of Finalism

B y the time o f the evol u tionary s y n thesis of the I 940s (see Chap­
ter 9) , virtu a l l y no evolutionary biologis t, in fa ct no com peten t
bio logi s t , was left who stil l believed in any final ca usat ion of cvo-

· 65 ·
 Th e Struggle against Ph ysicists and Ph ilosop hers

l ution . The few biologists who s till did so either had theologica l
com mit ments , l i k e Teilhard de Chardin , o r were unaware of the
developments of biolo gy in the twentieth cen tury, like Comte de
N o u y.
Final causes , h owever, are fa r more plausible and pleasing to a
layperson than the hap hazard and op portunistic process of na tu­
ral selection . For this reason , a belief in final cau ses has had a fa r
g reater hold outs ide of than within biology. A l n1ost all philoso­
p hers , for ins tance , who w rote on evolutionary ch ange in the one
hundred years a fter 1 8 59 were confirn1ed finalists , fro 111 Whe­
well , Herschel , and Mill to Henri B ergson in France, who pos­
tu lated a metap hysical force, ela n vital, which, even though he
disclai med its finalistic nature , could not have been anything ex­
ce pt a final cause, considering i ts effects . Whitehead , Polanyi , and
n1 any lesser p hilosop hers we re also fina listic . Throu ghout this
period there have been exceptions , the most noteworthy perhaps
bein g the Gern1an philosopher Sigwart, who as early as 1 8 8 1 pro­
vided a re n1 a rkably modern treatment of the problems of tele­
ology.
Modern philosophers-that is , those who have published since
the evolu tiona ry s ynthesis-have , on the whole, refrained from
invoking final causes when discus sing evolution ary progress .
A p parently they ful l y accept the explanation provided by the
evolutionary s ynthesis . When they do discuss teleology, l ike
Morton B eckner or Ernes t Nagel , they deal with adaptation and
\vi th " teleolo gical s yste n1 s . " Finalis n1 is no lon ger pa rt of any re­
spectable philosophy. One last v igorous a ttack on final is n1 wa s
Monad 's book Chance and Necessity ( 1 970) . But Monad fa iled to
unders tand the ex planatory power of natural selection and opted
fo r pure ch ance as having been responsible fo r the phenon1ena o f
natu re . S uch E picureanis m , ho\vever, is only ra rel y en countered
in n1 odern tirnes .
The rea son why the cont roversy about the validity o f teleolog­
ical thinkin g has been so indec isive has finally beco n1 e evident in
recen t years : the des igna tion u teleolog ical '' has been app lied to
fo ur quite differen t natural pheno1nena . Th ree of these can be ex-

· 66 ·
 The Struggle aga inst Ph y sicists and Ph ilosop h ers

plained b y science, while the fou rth one, an explanatory postu­

late for certain phenomena , has not been substantia ted .
( 1 ) Many seemin gly end-di rected processes or movemen ts in
inorganic natu re a re the simple consequences of natural laws . The
fal ling of a stone (due to gravity) or the cool ing of heated pieces
of n1ctal (owing to the second law of thermodyna n1 ics) arc cx­
an1ples of such t eleo,na tic processes , as such la w-di rected pro­
cesses are called.
(2) Processes in living organisms-as well as their behavior­
that owe their goal-directedness to the operation of an in born or
acquired program are called teleonotn ic. This includes all cha nges
in ontogenetic developmen t as well as end-directed behavioral ac­
tivities . Such processes can be analyzed st rictly scientifically,
since the end-point or goal is already contained in the program.
( 3 ) Adapted systems , like the heart, which pumps blood, or the
kidneys , which eliminate by-products of protein metabolism,
and which seem to work towa rd a goal have also been called te­
leological . An organ ism has hundreds , if not thousands , of such
adapted systems , from the molecular level up to the organism as
a whole, all of them acquired durin g the evolution of its ancestors
and contin uously fine-tu ned by natural selection . These systems
have the capacity for teleonomic behavior, but, being stationary,
are not themselves goal-seeking .
(4) From the Greeks on , there was a widespread belief that
everything in nature and its processes has a purpose, a predeter­
mined goal . And these processes would lead the world to ever­
greater perfection . Such a teleological worldview was held by
many of the great philosophers . Modern science, however, has
been un able to substantiate the ex istence of such a cosm ic teleology.
Nor have any mechanis n1s or laws been found tha t would permit
the funct ion ing of such a teleology. The conclus ion of science has
been tha t final causes of this type do not exist .
 C H AP T E R S I X
-------------• -------------

Darwin's Path to the Theor y

of Natural Selection

W HEN w E s P E A K of Da rwinism today, we mean evolution

b y natura l selection . The meaning of natural select ion , its
limits , and the processes b y which it achieves its effects are now
the most active areas of evolu tionary resea rch .
The fi fth one of Darw in's great evolutionary theo ries wa s his
most darin g , n1ost novel . I t dealt with the mechanism of evolu­
tionary chan ge and, more particularly, how this mechanism
could account for the seeming ha rmony and adaptation of the
organic world . It at tem pted to provide a natural explanation in
place o f the s upernatural one of natural theology. I n that res pect
Darwin's theory was unique; there was nothing like it in the
whole philosophical literature fro m the pre-Socratics to Des­
ca rtes , Leibniz, or K an t . It replaced teleolog y in nature with an
essen tially mechanical explanation .
To j udge fro n1 his writings , Darwin had a n1 uch sim p ler con­
cept of natura l selection than the modern evolutionist does . For
hi n1 there was a s teady produ ction of individual s , generation after
generation , some of whom were " su perior" in having a repro­
ductive advantage . For D a rwin selection was es sen tially a single­
step p roces s , the conveying of reprodu cti ve success . The n1 odern
evolutionis t agrees with Da rwin that the individual is the ta rget
of selectio n ; but we now also realize that natural selection is ac­
tua lly a two-s tep process , the first one consis ting of the produc­
tion of geneticall y different individuals (variation) , \vhile the sur­
vival and reproductive su cces s of these individuals is dctern1 ined
in the second step , the actual selection process . Although I have

· 68 ·
 Da rw in �� Path to the Theory of Natural Selection

called the theory of natural selection Darwin's fifth theory, It 1s

actually itself a small package of theories. This includes the
theory of the perpetual existence of a reproductive surplus, the
theory of a continuing availability of great genetic variability,
the theory of the heritability of individual differences, the theory
that mere reproductive superiority is selected for (sexual selec­
tion) , and several others.
The question concerning the conceptual sources of Darwin's
theory of natural selection is still highly controversial. A favorite
interpretation a mong historians has always been that it was a
manifestation of the thinking of upper-cl ass England in the first
half of the nineteenth century (consistent with empiricism, mer­
cantilism, industrial revolution, poor laws, and so forth) . l)ar­
win's admission that reading Malthus had given him the crucial
insight seemed to provide a powerful confirmation of this " exter­
nal causation. " The evolutionists, by contrast, have favored an
interpretation based on "internal causation," relying on Darwin's
insistence that his familiarity with the practices of the animal
breeders had provided him with the decisive evidence. The redis­
covery of Darwin's notebooks covering the one and a half years
prior to the date of his "conversion" has provided us with a great
amount of new information, but-although narrowing down
our options-it still permits conflicting interpretations. What I
present here is anything but the last word in a still-ongoing con­
troversy. It will require further research before the remaining dis­
agreements can be removed (Hodge and Kohn 1 9 8 5 ) .
Darwin had returned to England from the voyage of the Beagle
in October 1 8 3 6 . While working on his bird collections, and par­
ticularly through discussions with the ornithologist John Gould,
Darwin beca me an evolutionist, apparently in March I 8 3 7 (Sul­
loway 1 9 8 26) . Certainly by J uly 1 8 3 7 he had firn1 ly accepted evo­
lution by common descent. His new interpretation of the world
consisted not only in replacing a static or steady-state world by
an evolving one but also, and more important, in depriving n1an
of his uniq ue position ii) the universe and placing hin1 into the
stream of animal evolution. Darwin, after this date, never ques-
 D a rw in 's Path to th e Th eory of Na tura l Selection

tioned the fact of evolution, even though he continued for an­

other twenty years to collect supporting evidence. Yet, the causes
of evolution were at first a complete mystery to him.
For a year and a half Darwin speculated incessantly, developing
and then again rejecting one theory after the other (Kohn 197 5),
until he finall y had a decisive illumination on September 2 8 ,
1 8 3 8 . In his autobiography he describes it as follows (Darwin
1 9 5 8 : I 20) :

Fifteen months after I had begun my systen1atic enquiry, I hap­

pened to read for amusement Malthus on Population, and being
well prepared to appreciate the struggle for existence which every­
where goes on, fron1 long-continued observation of the habits of
animals and plants, it at once stru ck me that under these circun1-
stances favorable variations would tend to be preserved, and un­
favorable ones to be destroyed. The result of this would be the
forn1ation of new species. Here , then , I had at last got a theory by
which to work .

It was the theory later called by Darwin the theory of natural

selection. It was a most daring innovation, since it proposed to
explain by natural causes, mechanically, all the wonderful adap­
tations of living nature hitherto attributed to "design. "
Darwin makes it sound as though the concept of natural selec­
tion \Vas simplicity itself. But his memory deceived him. His
autobiography was written almost forty years later (in 1 8 76),
largely for the benefit of his grandchildren, and was replete with
characteristically Victorian self-denigrations. Darwin had forgot­
ten what a complex shift in four or five major concepts had been
required to arrive at the new theory. He probably never full y re­
alized himself how unprecedented his new concept was and how
totally opposed to n1any traditional assun1ptions.
Indeed, the concept of natural selection \Vas so strange to Dar­
win's contemporaries when he proposed it in the Or(gin o.f Sp ecies
that only a handful adopted it. Nearly three generations passed
before it becan1e universally accepted even a1nong biologists.
A111ong nonbiologists the idea is still unpopular, and even those

. 7 () .
 Da rt4 1 in �, Pa th to the Th eor y of Natu ra l Selection

who pay lip service to it often reveal by their comments that they
do not fully understand the working of natural selection. Only
when one is aware of the con1pletc unorthodoxy of this idea can
one appreciate Darwin's revolutionary intellectual achievement.
And this poses a powerful riddle: How could Darwin have arrived
at an idea which not only was totally at variance with the thinking
of his own tin1e but was so con1plex that even now, one and a half
centuries later, it is widely 1n isunderstood in spite of our vastly
greater understanding of the processes of variation and inheri­
tance?
Darwin's own version (in his autobiography) was that contem­
plation of the success of animal breeders in producing new breeds
had provided him with the clue for the mechanis1n of evolution
and was thus the basis for his theory of natural selection. We
know that this is a vast oversimplification-a revision of our
thinking which we owe to the rediscovery of Darwin's note­
books. In July 1 8 3 7 he had started to write down all the facts as
well as his own thoughts and speculations "which bore in any
way on the variation of animals and plants under domestication
and [ in] nature." Even though he later cut out occasional pages,
to use them for his book manuscripts, Darwin never discarded
these notebooks, and they were rediscovered in the 1950s among
the Darwin papers at the Cambridge University Library (Barrett
et al. 1987). Darwin's day-by-day records throw an entirely new
light on the development and the changes in his thought during
the period from July 1 87 3 to September 2 8, 1 83 8, when the
theory of natural selection was conceived.
One fact, the importance of which has not been reduced by the
recent discoveries, is the impact of Darwin's reading of Malthus.
The interpretation of the Malthus episode, however, has become
the subject of considerable controversy among the Darwin schol­
ars. According to some of them-de Beer ( 196 1) and S. Smith
( 1960), and, to a lesser extent, Gruber ( 1974) and myself-it was
merely the culmination in the gradual development of Darwin's
thinking, a little nudge that pushed Darwin across a threshold he
had already reached. According to others-Limoges ( 1970) and

. 71 •
 FIGURE I

Darwin's Explanatory M odel of Evolution through Natural Selection

Fact 1
Pote n t ia l expone n t ia l i n c rease
of pop u l a t io n s ( s upe rfec u n d i t y )
(S()fll7'f: Pa ley, M a l t h u s ,
a n d others)

Fact 2 I n ference 1
Observed stead,·-state St ruggle fo r e x i ste nce
s ta b i l ity of pop u l a t ions a m ong ind i \' id u a l s
(So11n-,: u n i versal (A111hor rl i1,frn'11ff:
obse rva t i o n s ) Malthus)

I n ference 2 I n ference 3
Fact 3 I ) i ffe re n t i J I sun i ,·a l . Thro ugh m ;,1 ny ge nera t io n s :
Li m i tat ion of resou rces i .e . n a t u ra l sel ect ion � e\'o l u t ion
Fact 4
(Soune: observat ion U n i q ue ne s s of t h e ( .4111hor ,f i1lnnu r: (A111hor ,f ll{fe't"t'llf f :
re i n forced by M a l t h u s ) indi, idual
I )am i n ) I h rn i n )
(So11nf: a n i m a l
b ree ders , taxo n o m i s t s )

Fact 5
H e r i t ab i l i t ,· of m u c h
o f t he i n d i , i d u a l
\'a r iat 1on
( Sount': a n i m a l breede rs)
 Daru, i11 's Path to the Theory o.f Natural Selection

Kohn ( 1980), for exan1ple-it constituted a rather drastic break,

almost equivalent to a religious conversion. W hich of these two
interpretations is nearer to the truth?
There are essentially two methods by which we can try to find
an answer. Either we can attempt to analyze all the entries in the
notebooks, in a chronological sequence, or we can try to recon­
struct Darwin's explandtory model of natural selection and then
study separately the history of each of its individual components.
My own choice is in favor of the second method, placed in a
chronological fran1ework, although both methods arc necessary
for a full understanding. The first method attempts to reconstruct
the trials and errors of Darwin's gradual approach, as reflected in
successive entries in the notebooks. It also exan1ines tentative
ideas that he later rejected. Gruber, Kohn, and, in part, Limoges
have favored this method.

D a rwin's Explanatory Model

What were the components of Darwin's explanatory model? For
n1y analysis I have found it most convenient to recognize five facts
and three inferences (see Figure 1 ). I shall attempt to determine,
first, at what time Darwin became aware of these five facts, then
at what time he made the three inferences, and whether or not
these inferences had already been made previously and could be
found in the literature.
The five facts were already widely known before the Malthus
episode, not only to Darwin but to his contemporaries, only a
single one of whom, A. R. Wallace, used them in exactly the
same way as Darwin. Merely having these facts obviously was
not enough. They had to be related to one another in a meaning­
ful manner; that is, they had to be placed in an appropriate con­
ceptual background. In other words, Darwin had to be intellec­
tually prepared to see the connections among these facts.
This leads us to the most interesting but also the most difficult
question: What had been going on in Darwin's mind in the one
and a half years prior to the Malthus episode? All the indications

. 73 ·
 D arw in 's Path to the Th eory of Natura l Selection

are that it was a period of u.nprecedented intellectual activity in

Darwin's life. Precisely Wfiat the changes in Darwin's thinking
were and how they were connected with one another has not yet
been investigated nearly as fully as it deserves. Gruber and Kohn
have examined this problem more carefully than anyone else, but
the Darwin correspondence of that period and other manuscript
materials that have not yet been thoroughly analyzed are bound
to provide new insights. My own tentative conclusions n1ay,
therefore, turn out to be incorrect. However, my reading sug­
gests to me that Darwin's beliefs chan ged moderately or drasti­
cally in four areas, which I shall simply list here and then discuss
in the context of Darwin's model.
( 1 ) Th e gradual rep lacetnen t o_( the assumption that all inditJidua ls o_(
a species are essen tia lly alike by th e concept of th e un iqueness o_( euery
indiv idual . As discussed previously, the belief that the observed
variability of phenomena reflects a limited number of constant,
discontinuous essences was gradually replaced by population
thinkin g, a belief in the reality of variation within a population
and in the in1portance of these individual differences. Most of
Darwin's earlier statements on species and varieties were strictly
typological. It is my in1pression that they became more popula­
tional as Darwin delved deeper into the literature of the anirnal
breeders and also later as a result of his work on the barnacles
(Ghiselin 1969) .
(2) A sh [{t _from so_(t to ward hard inh eritance . In his earlier state­
ments Darwin seen1ed to assume that most, if not all, inheritance
was "soft." He assumed that the material basis of inheritance is
not unchangeably constant but can be modified by use and dis­
use, by physiological activities of the body, by a direct influence
of the environn1ent on the genetic material, or by an inherent
tendency to progress toward perfection , and that these environ­
rnentally induced chan ges could be passed along to the next gen­
eration. This theory is called the inheritance of acquired charac­
ters. The growth of his population thinking, with its increasing
stress on genetic differences arnong individuals, indicates a gro\v­
ing awareness of the need to postulate "hard " inheritance-that

• 74 •
 Da rw in 's Pa th to the Th eory c�( Na tura l Selection

is , inheritance tha t is not a ffected directly by environ 1nental fa c­

tors .
(3) A changi11Jz attitude to 1vard the balance o_(nature. Da rwin began
to believe that the balance of nature is dyna1nic rather than static,
and he began to ask whether the balance is 1n ain taincd by benign
adj us t 1nen ts or by constan t war.
(4) A J:radual loss o_( his Christian _faith . Darwin lost his faith in
the years I 8 3 6 - 1 8 3 9 , n1uch of it clearly prior to the reading of
Malthus . In o rder not to hurt the feelings of his friends and of his
wife, Darwin often used deistic language in his pub lications , but
n1uch in his notebooks indicates that by this ti1nc he had become
a " materialis t " (more o r less equiva len t to an atheist; sec Chapter
2) .
These fou r changes in Da rwin's thinking are to so me extent
interconnected . Since they were largely unconscious, they are
usually reflected in D arwin's notebooks only in subtle chan ges of
wording , and there is con siderable leeway in possible interpreta­
tio n . However, keeping these four points in mind will sharpen
our awa reness of possible chan ges in Darwin's thinking in the
years p rior to reading Malthus, while we m ake a point-by-point
analysis of Darwin's explanatory model .

The S truggle for Exis tence

When recording his reaction to reading Malthus, Darwin makes
it quite clear that it was not M althus 's general attitude that had
acted as a catalyst on his thou ghts but one pa rticular sentence, for
he says that " yet until the one sentence of Malthus no-one clea rly
perceived the g reat check a m ongst men " (D : 1 3 5 ) . De Beer
( 1 963 :99) su cceeded in determinin g what Malthus 's crucial sen­
tence was : " I t m ay safely be pronounced , therefore, that the pop­
ulation , when unchecked , goes on doubling itself every twenty­
five yea rs , or increases in a geometrical ratio . " From then on ,
Darwin stressed that it was M althus's demons tra tion of the ex­
ponen tial in crease of popula tion s that was decisive in his discov­
ery o f the im po rtance of natural selection (Fact I) .

• 75 •
 Da rw in 's Path to the Theory of Natural Selection

Yet there is a puzzling difficulty. W hy did it take Darwin so

long to recognize the evoi�tionary significance of the Malthusian
principle? The prodigious fertility of animals and plants had been
pointed out by many of Darwin's favorite and most frequently
read authors, like Erasmus Darwin, Charles Lyell, Alexander von
Humboldt, and William Paley. Furthermore, Malthus's principle
was widely discussed in the essay literature of the period. Why
then did this suddenly impress Darwin so profoundly on Sep­
tember 2 8 , 1 8 3 8 ?
Four reasons might be suggested-the first one being , as
pointed out by Gruber ( 1 974) , that Darwin had learned on the
three preceding days (between September 25 and 27) of the un­
believable fertility of protozoans by reading Ehrenberg 's work on
the subject. This quite likely primed Darwin's receptivity for
Malthus's thesis. The second reason is that when Malthus applied
the principle to man, a species with relatively few offspring, Dar­
win suddenly realized that a potentially exponential increase of a
population was entirely independent of the actual number of off­
spring of a given pair. The third reason is that the Malthus epi­
sode came at a time when population thinking had begun to ma­
ture in Darwin's mind. The fourth reason, suggested by Ruse
( 1 979 : I 75) , is that the numerical formulation sug gested by Mal­
thus seemed to satisfy the mathematical requirements of such
Newtonians as Herschel.
The second fact in Figure I -population stability-was not in
the slightest bit controversial. No one questioned that the num­
ber of species and, aside from ten1porary fluctuations, the num­
ber of individuals in every species maintained a steady -state sta­
bility. This is implicit in the concept of plenitude of the
Leibnizians and in the harn1ony-of-nature concept of the natural
theologians. If there is an extinction, it is balanced by speciation,
and if there is high fertility it must be counterbalanced by n1ortal­
ity. In the end everything adds up to a steady-state stability.
The third fact-lin1itation of resources-again was not at all
controversial, being very n1uch part of the balance-of-nature con-

· 76 ·
 Da rw in 's Pa th to the Th eo ry of Na tural Selection

cept of natural theology, so dominant in England in the first half

of the nineteenth century.
Darwin's first great inference, derived from these three facts,
was that exponential population growth combined with a fixed
an1ount of resources would result in a fierce struggle for exis­
tence . We must ask if this inference was original with Darwin,
and if so, what part of it did he owe to Malthus? This is perhaps
the most controversial question raised by the analysis of the selec­
tion theory. The main difficulty is that the term "struggle for ex­
istence " and similar synonymous terms were used in different
senses b y different authors.
Before we can analyze then1 , we must deal with one other con­
cept, the idea of a perfect balance of nature, an idea prevalent in
the eighteenth century : nothing in nature is too much, nothing
too little, everything is designed to fit with everything else . Rab­
bits and hares have lots of young b � cause food must be available
for foxes and other carnivores. The whole economy of nature
forms a harmonious whole that can in no way be disturbed. This
is why Lamarck, who was very much an adherent of this concept,
could not conceive of extinction. Cuvier likewise had adopted the
idea of perfect balance , as shown in correspondence with his
friend Pfaff. He transferred the same concept to the structure of
an organism, which he visualized as a "harmonious type " in
which nothing cou ld be changed. Everything in such a complex
system is so perfect that any change would lead to deterioration.
This type of thinking was still dominant in Darwin's day, not
only among the natural theologians of England but also on the
Continent. Indeed, one can find a number of entries in Darwin's
notebooks that seem to reflect this kind of thinking. But was
Darwin still a wholehearted supporter of the concept of a har­
monious balance of a benign nature? This is a very important
question because it affects the interpretation of what Darwin
understood under the term "struggle for existence. "
For u s moderns the term means a fierce fight with no holds
barred. But for the natural theologians the struggle for existence

· 77 .
 Darwin 's Path to the Th eory of Na tu ral Selection

was a beneficial feedback device, the function of which was to

maintain the balance of na.ture. It is, as Herder ( 1 7 84) called it,
"the balance of forces which brings peace to the creation." Lin­
naeus ( 1 7 8 1 ) devoted an entire essay to the " Police of Nature" and
emphasized that "those laws of nature by which the number of
species in the natural kingdoms is preserved undestroyed, and
their relative proportions kept in proper bounds are objects ex­
tremely worthy of our attentive pursuit and researches." Lamarck
expressed similar sentiments.
Was this benign interpretation of the struggle for existence
unanimous? Unfortunately, even today we have no reliable anal­
ysis that would give us an answer to this question. My in1pression
is, however, that as the interaction of predators and prey, of
parasites and their victin1s, the frequency of extinction, and
the struggles of con1peting species became better known, the
struggle for existence was more and more recognized as a "war"
or fight, a struggle for survival, "red in tooth and claw" as Ten­
nyson later expressed it. Bonnet ( 1 78 1 ) and de Candolle ( 1 820)
en1phasized that this war among species consisted not n1erely of
predator-prey relationships but of competition for any and all
resources. However, it was not at all appreciated how fierce this
struggle is, and Darwin admits that "even the energetic language
of de Candolle does not convey the warring of the species as [ con­
vincingly as does the] inference from Malthus."
Nevertheless, it is highly probable that Darwin had been grad­
ually conditioned by his reading to a far less benign interpretation
of the struggle for existence than that held by the natural theolo­
gians. The mere fact that Darwin had adopted evolution 1nust
have 1nade him aware of the frequency of extinction and of the
unbalances and adaptational lags caused by evolutionary changes.
Fro1n Aristotle to the natural theologians it was considered axio­
matic that a belief in a harmonious universe and perfect adapta­
tion in nature, or in a Creator continuously active in correcting
in1perfections and imbalances, \Vas incon1patible with a belief in
evolution. By necessity, accepting evolutionary thinking under-
 Da rwin s Pa tlz to th e Th eor y of Na tura l Select ion

mined a continued adherence to a belief in a harmonious uni­

verse.

St ruggle an1on g Species or Individuals ?

Of far greater in1portance is a second question: Between whon1
does the struggle for existence take place? This question allows
two drastically different answers . In the entire essentialistic liter­
ature the struggle is considered to take place a1nong species . The
balance of nature is maintained by this struggle, even if it occa­
sionally causes the extinction of a species . This is the interpreta­
tion of the struggle for existence in the literature of natural the­
ology, up to de Candolle and Lyell, and is the major e1nphasis of
Darwin's notebooks up to the Malthus reading . The main func­
tion of this struggle is to correct disturbances in the balance of
nature, but it can never lead to changes ; on the contrary, it is a
device to preserve a steady-state condition. As such it continued
even after 1 8 3 8 to be an important component of Darwin's think­
ing , particularly in his biogeographic discussions (such as deter­
mination of species borders) .
Only when one applies population thinking to the struggle for
existence can one make the crucial conceptual shift to recognizing
a struggle for existence among individuals of a single population.
This , as Herbert ( 1 97 1 ) was the first to recognize clearly, was
Darwin's decisive new insight resulting from his reading of Mal­
thus , although Mayr ( 1 9 5 96) and Ghiselin ( 1 969) had previously
pointed out the populational nature of selection. If most individ­
uals of every species are unsuccessful in every generation, then
there must be a colossal competitive strug gle for existence among
them. It was this conclusion that made Darwin think at once of
various other facts that had been slumbering in his subconscious
b ut for which, up to that moment, he had had no use.
Darwin's reading of Malthus was dramatic and climactic, and
it does not matter whether one interprets it as a con1plete reversal
of Darwin's thinking or whether one believes that "the evidence

· 79 ·
 D a rw in 's Pa th to th e Theory o_f Natural Selection

suggests that the change in �hoice of unit was a protracted pro­

cess, stretched over a year ·o r more, and linked to other aspects of
his thought" (Gruber 1974). I n1yself hold with the latter view,
because the capacity to be able to interpret the Malthus staten1ent
on exponential growth of populations and to apply it to individ­
uals requires population thinking, and this Darwin had been
gradually acquiring during the preceding year and a half . That
every thing came to a dramatic climax on September 2 8, 1 83 8,
however, is beyond question.
The whole concep t of competition among individuals would
be irrelevant if all these individuals were ty pologically identical­
if they all had the same essence. Competition does not become
meaningful in an evolutionary sense until a concept has devel­
oped that allows for variability among the individuals of the san1e
population. Each individual may differ in the ability to tolerate
climate, to find food and a place in which to live, to find a n1ate,
and to raise young successfully. The recognition of the unique­
ness of every individual and the role of individuality in evolution
is not only of the utmost importance for an understanding of the
history of biology, but it is one of the most drastic conceptual
revolutions in Western thought (Fact 4).
There is little doubt that this concept-which we call "popu­
lation thinking"-received an enormous boost in Darwin's mind
through his reading Malthus at that right moment. Yet, curi­
ously, when we go through Malthus's writings we find no trace
of population thinking. There is nothing whatsoever even faintly
relating to the subject in those early chapters of Malthus that gave
Darwin the idea of exponential growth. There is a reference to
animal breeding in chap ter 9, but there the subject is introduced
to prove exactly the opposite point. After referring to the claims
of the animal breeders, Malthus states, "I am told that it is a
n1axim among the in1provers of cattle that you may breed to any
degree of nicety you please, and they found this maxim upon
another, ,vhich is, that some of the offspring will possess the de­
sirable qualities of the parents in a greater degree." He then pro-

· 8o •
 J. S . Henslow ( 1 796 - 1 8 6 1 ) , Darwin's bo tany
professo r at C a n1bridge University, who pro­
cured for Darwin the invitation to join the
Beagle voyage

The H. M . S . Beagle, in the Straits of Magellan , 1 8 3 3

 To Cape Verdes,
Azores and Falmouch
< Z On urn,)

From I >cvonporl
o_
IP ( .? 7 Del· UB I )
nc> (1§ ·'
C)�t
From Asccnsmn I sland
(see large map)

oquimbo
Santa Fe
Valparaiso � / ��e nd oza T
Mercedes
'. \ Montevideo
,\ tp.,
iago Buenos, Maldonado
Aires /
Western lsl ;.v /\
.�,-�
'.'\OJffH
l '.-\CI FIC NORTI I ATLA ,TIC OCEAN
< >< :EA'.'\ The course of the;Bfagle
and Darwin's inlancfjourneys
in South America

" DI AN Santa Cr 1 1 , 'Ht . � �Jil:1.- Falkland Is.

(;alapagos OCEAN River
. /,
.� · \brquna, Tierra del

c:.
!\·1adagas celing ls� .:> ·
---:d•! • �1,
Sol i<. lslanJ,
' l ahi1i uritius
Bourbon Is.
Valpa /of
/ ��!ncds

<. :ape: of
S< >l Tl I PA< :1 Fl< : < >< :1•:..\, ( ;,,oJ :i� ,r
I lope
S< >F rt l :ffL:\ NTIC OCEA!\

:alkland Is.

:< :apcl lorn

Straits of :\1agcllan The Voyage of the Beagle ( 1 83 1-6)

The route of the voyage of the Beag le, 1 8 3 1-1 8 3 6

The naturalist Jean Baptiste
L a marck ( 1 744- 1 8 29) , the fi rst
person to adopt a consistent
theory of gradual e volutionary
change
Charles Lyell ( 1 797- 1 8 7 5 ) ,
geologist, to whose theory of
uniformitarianis m Darwin
owed much of h is thinking
about evolution ary change

The o rnithologist John Gould ( 1 804- 1 8 8 1 ) ,

who info r med Darwin in March 1 8 3 7 that the
specimens of mockingbirds he had collected on
th ree islands in the Gala pa gos were th ree dis­
tin ct species
Em1na Wedgwood Darwin ( 1 8 0 8 - 1 896) in 1 840, Charles Darwin's wife
and first cousin, and daugh ter of the famous potter Josiah Wedgwood I I
Charles Darwin in 1 840
The botanist Joseph Dalton Hooker ( 1 8 1 7-1 9 1 1 ) ,
Darwin's friend and supporter, who i n 1 8 5 8 , with
Lyell , presented Darwin's and Wallace's findings to
the Linnean Society o f London

The morphologist and paleontologist

Richa rd Owen ( 1 8 04- 1 892) . once Darwin's
friend . who viciously attacked the Or(gi11 and
whose s taten1er1ts Hu xley attacked \Vith
equal vigor
 Alfred Russel Wal lace ( 1 8 2 3 - 1 9 1 3 ) , co-discoverer with
Darwin of the theo ry of evolu tion through natural se­
lection and opponen t , with Wei s rnann, of the theo ry of
the inheritance of acquired characters

T h e 111 o r p h o I o g i s t , p h y s i o I o g i s t , a n d e 111 b r yo I o g i s t
T h o m a s Hen r y H u x l e y ( r 8 2 5 - 1 8 9 5 ) , sel f-a n o i n t e d a s
" D a r w i n 's b u l l d o g " fo r h i s e n e r g e t i c d e fe n s e o f D a r w i n
a n d t h e t h e o r y o f d e s ce n t
.A.bo 11e: The botan ist Asa Gray ( 1 8 1 0 -
1 8 8 8 ) , Darwin's mos t importan t sup­
porter in America and a devout C h ris­
tian who su cceeded in reconciling na tu­
ral selection \V ith a belief in a personal
god

Belo w : The S w i ss-bo rn naturalist Louis

Agassiz ( 1 809 - 1 8 7 3 ) , an ou tstanding
ich thyologist and specialist of o ther
groups of organis m s , who p layed a
pron1i nen t role in the spread of natural
historv studies in A1nerica : a devout cre­
ationist. A gassiz cha racterized the Orig i11
as ua scien tific 1n istake. untrue in its
fact s . un scien t ific in its n1ethod . and
111 ischievous in its ten den cies "
A ho ve: The b i o l o g i s t E rn s t Ha eckel ( 1 8 3 4- 1 9 1 9) , an
en t h u s i a s t i c s u ppo rter and popu la rizer o f D a rw i n i s n1
in Ge r m a n y, w h o spec i a l l y e m phasized the s t u d y of
p h y l ogen y

B cl o u , : A u gus t Wei s 111 an n ( 1 8 3 4 - I 9 I 4) , th c n i n e t cc 11 th

cen tu ry 's g rea tes t e vo l u t i o n i s t a fte r D a r win and a
st a u n c h s u pporter o f n a t u ra l selec tion ; he was res pon­
si b l e fo r t h e d e m i se o f t h e t h eory o f acq u i red c h a rac­
ters a n d l a i d t h e fou n d a t i o n s o f gen e t i cs
C h a rles Darwin on the veranda a t Down House
 Darwin �� Pa th to the Theory o_f Natural Selection

duces all sorts of facts and reasons why this cannot possibly be
the case, leading him to the conclusion that "it cannot be true,
therefore, that an1ong animals, some of the offspring will possess
the desirable qualities of the parents in a greater degree; or that
animals are indefinitely affectable" (Malthus 1 79 8 : 1 6 3 ) .
Where, then, did Darwin get his population thinking, since he
evidently did not get it fron1 Malthus? In his autobiography and
in various letters Darwin e1nphasized again and again that he had
been mentally prepared for the Malthus principle by studying the
literature of animal breeding. Recent commentators such as Lim­
oges and Herbert have insisted that this must be a lapse of Dar­
win's memory because there is very little about animal breeding
in Darwin's notebooks until about three months after the Mal­
thus reading. For myself, I am convinced that Darwin's own pre­
sentation is nevertheless essentially correct.
If we ask ourselves what Darwin would be likely to enter in his
notebooks, we would certainly say new facts or new ideas.
Hence, since it was not a new subject, animal breeding surely
wou ld not qualify. Darwin's best friends at C ambridge Univer­
sity were the sons of country squires and of owners of estates.
They were the " horsy set," riding or hunting with dogs on every
occasion (Him melfarb 1 9 5 9) . All of them to a greater or lesser
degree were interested in animal breeding. They must have ar­
gued a great deal among themselves about Bakewell and Sebright
and the best methods of breeding and improving dogs, horses,
and livestock.
How else-other than that it had a great interest for him-can
one explain that Darwin, in the excessively busy period after the
return of the Beag le, devoted so much of his time to studying the
literature of the animal breeders? To be sure, Darwin's primary
interest was in the origin of variation, but in the course of his
reading Darwin could not help absorbing the important lesson
from the breeders-that every individual in the herd was differ­
ent from every other one and that extreme care had to be used in
selecting the sires and dams from which to breed the next gener-

. 81 •
 Darw in 's Path to the Theory of Natu ral Selection

ation. I am quite convinced that it is no coincidence that Darwin

studied the literature of th€. 'animal breeders so assiduously ex­
actly during the six months before reading Malthus (Ruse 1 97 5 a) .
It was not the process of selection but the fact of the differences
among individuals that Darwin remembered when suddenly be­
com ing aware of the competition among individuals, of the
struggle for existence among individuals. Here we have the for­
tuitous com ing together of two important concepts -excessive
fertility and individuality-which jointly provide the basis for an
entirely new conceptualization.
Variation can be of evolutiona ry significance-that is, it can be
selected-only if at least part of it is heritable (Fact 5 ) . Like the
animal breeders from whom he got so much of his inforn1 ation,
Da rwin took this heritability of characters con1pletely for
granted. And this assumption can be held quite independently of
one's assumptions concerning the nature of the genetic material
and of the origin of new genetic factors . Darwin's ideas on these
subjects were quite confused, but he did know a number of things
by observation. He knew that in asexual reproduction the off­
spring are identical with the pa rent, while in sexual reproduction
offspring are different from the parent and from one another.
Furthermore, he knew that each offspring had a n1ixtu re of the
cha racters of both parents . On the whole Darwin treated genetic
variation as a " black box . " As a naturalist and reader of the animal
breeding literature, he knew that variation was always present,
and this is all he had to know. He was also convinced that the
supply of variation was renewed in every generation and thus was
always abundantly available as raw material for natural selection.
In other words , a correct theory of genetics was not a prerequisite
for the theory of natu ral selection.

The Path to Discovery

The next question we have to answer is how Darwin arrived at
the actual concept of natural selection on the basis of the stated
five facts and his first inference . In his autobiography ( 1 9 5 8 : 1 1 8 -

· 82 ·
 Darwin 's Pa th to th e Theory of Natur(l/ Selection

1 20) Darwin stressed that he " collected facts on a wholesale scale ,

more especially with respect to domesticated productions, by
printed enquiries, by conversation with skillful breeders and gar­
deners, and by reading . . . I soon perceived that selection was
the key-stone of rnan's success in making useful races of animals
and plants. But how selection could be applied to organisn1s liv­
ing in a state of nature ren1ained for some time a mystery to me ."
In I 8 5 9 he wrote to Wallace, "I came to the conclusion that selec­
tion was the principle of change fron1 the study of domestic pro­
ductions; and then, reading Malthus, I saw at once how to apply
this principle ." To Lyell he wrote, with reference to Wallace's
theory, " We differ only [in] that I was led to my views from what
artificial selection had done for domestic anin1als." Traditionally,
these statements were accepted by Darwin students as a correct
representation of the facts .
This interpretation of Darwin's path to the concept of natural
selection has been challenged in recent years in the wake of the
discovery of Darwin's notebooks, for the same reason that re­
searchers have come to doubt the sources of Darwin's population
thinking. Limoges and Herbert point out that in the first three
notebooks Darwin nowhere refers to selection or to the selective
activities of animal breeders , particularly in the production of
new domestic races . They claim that Darwin was interested in
domestic animals only because he hoped to find evidence con­
cerning the occurrence of variations and the mechanisms of their
production, matters that are difficult to study in wild popula­
tions.
It is true that the term "selection" does not occur in Darwin's
notebooks; it is first found in his 1 842 sketch in the words "natu­
ral means of selection" ( F. Darwin 1 909: 1 7) . Darwin here refers
to artificial selection by the term "human selection. " Actually, in
the notebooks Darwin not infrequently refers to the process of
selecting, but he uses a different term- "picking."
I am willing to grant to the recent critics that there is no evi­
dence in the notebooks of a simple application to the evolutionary
process of the analogy between selection by n1 an and selection by

• 83 .
 Da rwin s Pa th to th e Theory of Natural Selection

nature. This is quite evident when one reads the crucial entry in
the notebooks on Septemk,er 2 8, 1 8 3 8 (here reproduced in the
original telegraph style):

Take Europe on an average every species must have san1e number

killed year with year by hawks, by cold etc. -even one species of
hawk decreasing in number must affect instantaneously all the
rest. -The final cause of all this wedging, must be to sort out
proper structure, and adapt it to changes-to do that for form,
which Malthus shows is the final effect (by means however of vo­
lition) of this populousness on the energy of 1nan. One may say
there is a force like a h undred thousand wedges trying [ to ] force
every kind of adapted structure into the gaps in the economy of
nature, or rather forming gaps by thrusting out the weaker ones.

The metaphor here is "wedging," not "selecting." Thus it ap­

pears that the arguments of the critics have considerable validity.
However, the analogy bet ween artificial selection and natural se­
lection is not necessary for Darwin's conclusions. Inferen ce I and
Fact 4 automatically result in Inference 2 (natural selection). I t is
quite likely that Darwin did not see the obvious analogy between
artificial and natural selection until some time after the Malthus
reading. Yet, I have little doubt that the copious reading Darwin
had done in the field of animal breeding had prepared his mind to
appreciate the role of the individual and its heritable qualities.
Indeed, I am convinced, with Ruse, that the many years during
which Darwin had been exposed to the ideas of the animal breed­
ers had preconditioned his mind to appreciate the importance of
the Malthus principle. This dormant knowledge was actualized
under the impact of reading Malthus.
The natural selection of individuals with particular heritable
qualities, continued over many generations, automatically leads
to evolution, as in Inference 3 . In fact, this process is sometimes
used as the definition of evolution. In this connection it n1ust be
en1phasized once more that Darwin's inference is exactly the op­
posite of that of Malthus, who had denied that "some of the off­
spring will possess the desirable qualities of the parents in a

• 84 .
 Da rwin s Path to the Theory of Natu ral Selection

greater degree . " Indeed , Malthus used his entire argu men t as a
refu ta tion of the thesis o f C ondorcet and Godwin of human per­
fectibility. The M althusian principle, dealing with populations of
essentialistically iden tical individuals , causes only quantitative,
not q ualitati ve, changes in populations (Limoges 1 970) . The fre­
quentl y u pheld thesis that it was the social-sciences message of
M althus that was res ponsible for Darwin's new insight has been
convincingly refuted by Gordon ( 1 989) .

How Great Was D a rwin's Debt to M althus?

That t h e M althus reading a cted a s a catalyst i n Da rwin's mind i n
producing the theory o f natu ral selection cannot b e disputed and
was emphasized by D a rwin himself again an d again . However,
when we analyze the co m ponents of the theory, as we have j ust
done, we find that it is p rim arily the insight that com petition is
among individuals rather than species that is clearly a Malthusian
contribu tion . To be sure , this in turn· led Darwin to a reevaluation
of other phenomena , such as the nature of the struggle for exis­
tence, but only as second-order consequences . I agree with those
who think that the M althusian thesis of exponential growth was
the capstone o f D arwin's theory. "The one sentence of Malthus "
acted like a crys tal dro pped into a su percooled fluid.
There i s , however, also a second and more subtle Malthusian
impact . The world o f the natural theologians was an optimistic
world : everything that was happening was for the common good
and helped to maintain the perfect harmony of the world. The
world o f M althus was a pessimistic world : there are ever-repeated
catastrophes , an unending, fierce struggle for existence, yet the
world es sentially remains the same. However much Darwin
might have begun to question the benign nature of the struggle
for exis tence, he clea rly did not appreciate the fierceness of this
stru ggle before reading Malthus . And it permitted him to com­
bine the bes t elements of Malthus and of natural theolog y : it
brought him to the belief that the struggle fo r existence is not a
hopeless s teady-state condition, as Malthus believed, but the very

• 85 •
 Da rw in 's Path to the Theory of Natura l Selection

means by which the harmony of the world is achieved and main­

tained. Adaptation is the res.ult of the struggle for existence .
The events of September 2 8 , 1 8 3 8 , are of great interes t to stu­
dents of theory formation. Given the ex tent to which Darwin
was in possession of all the other pieces of his theory prior to this
date, it becomes clear that in the case of a complexly structured
theory it is not sufficient to have most of the pieces ; one n1ust
have them all . Even a small deficiency, like defining the word
"variety" typologically ins tead of populationally, rn ight be suffi­
cient to prevent the correct piecing together of the con1ponents .
Equally important is the general ideological attitude of the
theory-builder. A person like Edward Blyth n1ight have had in
his possession the very same co1nponents of the theory as Darwin
but would have been unable to piece them together correctly ow­
ing to incompatible ideological comn1itments . Nothing illus­
trates better how important the general attitude and conceptual
framework of the maker of a theory is than the sirnultaneous,
independent proposal of the theory of natu ral selection by A. R.
Wallace . He was one of the few people, perhaps the only one,
who had had a similar set of past experiences: a life dedicated to
natural his tory, years of collecting on tropical islands , and the ex­
perience of reading M althus .

What Is Natural Selection ?

Darwin's choice of the word "selection" was not particularly for­
tunate . It suggests some agent in nature who, being able to pre­
dict the future, selects " the best." This , of course, is not what
natural selection does . The term simply refers to the fact that only
a few (on the average, two) of all the offspring of a set of parents
su rvive long enough to reproduce . There is no particular selective
force in nature, nor a definite selecting agent . There are many
possible causes for the success of the few survivors . Some sur­
vival, perhaps a lot of it, is due to stochastic processes , that is,
luck. Mos t of it, though, is due to a superior working of the
physiology of the surviving individual, which permits it to cope

· 86 ·
 Da rw in �, Pa th to th e Th eory of Na tu �al Selection

with the vicissitudes of the environn1ent better than other n1en1-

bers of the population. Selection cannot be dissected into an in­
ternal and an external portion. What detennines the success of an
individual is precisely the ability of the internal n1achincry of the
organism's body (including its in1n1une syste111) to cope with the
challenges of the environ1nent. It is not the environn1ent that se­
lects, but the organis111 that copes with the environ1nent n1ore or
less successfully. There is no external selection force.
A few examples 1nay illustrate this. Let us take, for instance,
resistance against pathogens. Bacteria and other pathogens rep­
resent the environn1ent; an anirnal's defense against then1 consists
of intracellular selection processes. Likewise, adaptation to the
temperature of the environment is controlled by balanced physi­
ological mechanisn1s, regulated by feedback mechanisn1s. The
success of an organism depends to a great deal on its normal de­
velopment fron1 the fertilized egg to adulthood. Almost all de­
partures from normalcy in development will be selected against.
Considering that for many people the term "selection" has a
teleological connotation-that is, it suggests purpose-many al­
ternative terms have been proposed, such as "survival of the fit­
test," "selective retention," "biased nonelimination," and so
forth. W hat all these terms try to n1ake clear is that selection is an
a posteriori phenomenon-that is, it ·is the survival of a few indi­
viduals who are either luckier than the other members of the pop­
ulation or who have certain attributes that give them superiority
in the particular context. The probabilistic nature of selection
cannot be stressed too strongly. It is not a deterministic process. ·1
\
Moreover, because selection is a very broad principle, it is prob- � , l · '

ably not refutable (Tuomi 1 9 8 1 ). However, each concrete appli- !

cation of the principle of natural selection to a specific situation is
testable and refutable.
One must distinguish between two applications of selection.
" Selection of " specifies the target of selection, and this is nor­
mally (in sexually reproducing organisn1s) a potentially repro­
ducing individual, as represented by its phenotype (body). For
this reason it is confusing to say that the gene is the unit of selcc-
 Darwin 's Path to the Th eory of Na tu ral Selection

tion. " Selection for" specifies. ' the particular phenotypic attribute
and corresponding componait of the genotype (DNA) that is re-
sponsible for the su ccess of the selected individual . The now­
obsolete concept that evolution is the interplay between genetic
mutation and selection was part of the saltationist thinking of the
Mendelians , as we will see. The material with which selection
/ works is not mutation but is rather the recombination of pa rental
) genes , which produces the new genotypes that direct the devel­
opment of individu als which are then exposed to selection in the
next generation. It must always be remembered that selection is a
two-step process . The first step consists in the production
(through genetic recombination) of an immense amount of new
genetic variation, while the second step is the nonrandom reten­
tion (survival) of a few of the new genetic va riants .
Selection at the level of the whole organis m results in changes
at two other levels: that of,the gene, where through the selection
of individuals certain genes may increase or decrease in frequency
in the population, and at the level of the species , where the selec­
tive superiority of members of one species may lead to the extinc­
tion of another species . This process , as mentioned earlier, has
often been called species selection but is perhaps better called spe­
cies replacement or species succession, in order to avoid misinter­
pretations . (Nothing is ever selected "for the good of the spe­
cies .")
Finally, it must be pointed out that two kinds of qualities are at
a pren1ium in selection. What Darwin called "natu ral selection"
refers to any attribute that favors survival, such as a better use of
resources , a better adaptation to weather and climate, superior
resistance to diseases , and a greater ability to es cape enemies .
However, an individual may make a higher geneti c contribution
to the next generation not by having superior survival attributes
but merely by being more successful in reproduction. Da rwin
called this kind of selection "sexual selection. " He was particu­
larly in1pressed by male secondary sexual characters , such as the
gorgeous plumes of n1ale birds of paradise, the gigantic size of
bull elephant seals , or the impressive antlers of stags . Modern

· 88 ·
 Da rwin 's Path to the Theory of Na tu_,ral Selection

research has shown that selection favors their evolution either be­
cause they aid in con1petition with other males over access to fe­
males , or because females are attracted to mates with these char­
acteristics . This latter process is known as "female choice ."
Selection for reproductive success affects many life history traits
beyond sexual dimorphism.
The path by which the theory of evolution by natural selection
was gradually clarified and modified will be described in the fol­
lowing chapters . Eventually the theory was universally adopted
among biologists , a developtn ent I refer to as the second Darv1in­
ian revolution.
 C H A P T. "E R S E V E N
-------------• -------------

What Is Darwinism?

C H AR LE s D AR w I N was the most talked about person o f the

I 8 6os . T. H. H u xley, always a coiner o f felicitous ph rases ,
soon referred to D arwin's ideas as " Darwin isrn " ( 1 864) , and in
1 8 89 A l fred Russel Wallace published a whole volun1e en titled
Darwin ism . However, since the 1 86os no two authors have used
the wo rd " D arwinis1n " i n exactl y the san1e way. As in the old
story of the three blin d n1en and the elephan t, every writer on
Darwinisn1 seemed to touch upon on ly one of the n1 any as pects
of D arwin is m , all the while thinking that he had the real essence
of what this term signifies . Thu s , everybody who read the Origin
responded onl y to those parts o f it that ei ther sup ported his own
precon ceived ideas or were i n conflict with them . What these
w riters failed to gras p is that Darwinis n1 is not a monolithic
theory that rises o r falls depending on the va lidi t y or invalidi ty of
a single idea .
This n1onolithic tradi tion actually s tarted with Darwin him­
self, who often s poke o f his " theory o f descen t with modification
through n a tu ral selection " ( 1 8 5 9 : 4 5 9) as though the theory of
co mmon descent was inseparable from that of natural selecti on .
How separable the t wo theo ries actually were was demons trated
when aln1ost every knowledgeable biologist adop ted the theo ry
of co n1 mon descent soon a fter 1 8 5 9 but rej ected natural selection .
They explained descent instead b y Lamarckian , finalis t, o r sa lta­
tional theories (Bowler 1 9 8 8 ) . A n u m ber of passages in the On�(!in
indicate how confused Darwin him self was on the subj ect : " The

. 90 •
 What Is Darw in ism ?

fact, as we have seen, that all past and present organic beings con­
stitute one grand natural system, with grou p subordinate to
grou p , and with extinct groups often falling inbetween recent
grou ps, is intelligible on the theory of natu ral selection"
( I 8 5 9 : 4 78) . Actually, the hierarchical organization of the living
world is ex plicable by the theory of con1mon descent, but this
tells us absolutely nothing about the mechanism by which these
changes were effected.
Even in modern tin1es far n1ore authors speak of Darwin's
theory in the singular than acknowledge the heterogeneity of
Darwin's paradigm. Even authors like Kitcher ( I 9 8 5) and Burian
( 1 9 8 9) , who are aware of the complexity of Darwin's paradigm,
continue to refer to Darwin's theory in the singular. Bu rian calls
the synthetic theory of evolution "the current variant of Darwin's
th eory. "
How Darwinism is seen depends to a large extent on the back­
ground and the interests of the viewer. The word has a different
meaning for a theologian, a Lamarckian, a Mendelian, or a post­
synthesis evolutionary biologist. Another dimension that con­
tributes to the diversity of opinion about the meaning of Darwin­
ism is geography: the word "Darwinism" has meant something
different in England, in Germany, in Russia, and in France. From
the beginning, as we have seen, Darwin's theories were in oppo­
sition to a number of ideologies such as essentialism, physicalism,
natural theology, and finalism whose strength varied fron1 one
country to the next. For the su pporters of one or the other of
these ideologies, the word "Darwinism" stood for the opposite
of their own beliefs.
An equally great diversity exists in the time dimension. Con­
cepts differ from facts by continuing to change over time. Hull
( 1 9 8 5 ) has rightly referred to "conceptual development as a gen­
uinely temporal process in which real change occurs. " What was
called Darwinism in 1 8 5 9 was no longer considered so thirty
years later, because the term had been transferred to something
very different from that which it designated at the earlier period.

• 91 .
 What Is Darwinism ?

This fact was well stated by Wallace in the preface to Da rwinisn1

( 1 8 89) . Here he explained .that Darwin had done such an excellent
j ob in proving descent with modification that this theory was
now universally accepted as the order of nature in the organic
world . " The obj ections now made to Darwin's theory apply,
solely, to the particular means by which the change of species has
been brought about, not to the fact of that change ." A las , Wallace
was way ahead of his ti me in his cham pionship of natural selec­
tion .
Differen t co mponents of Darwin's paradigm were particularly
interesting at different periods . At each stage in the history of
D arwinism a differen t one of Darwin's theories was referred to as
Darwinis m : anticreationis m vs . Christian orthodoxy, gradualism
vs . Mendelian saltationis m , selectionism vs. Lamarckis m or fi­
nalism , and so on . This continuing change of meaning poses the
awkward question of what establishes the continuity among all
these Darwinis m s ? Do these various Darwinisms have anything
in co m mon? The an swer of course is that they are all based on
Darwin's original paradigm, as presented in the Origin .
The best way to document the great variety of meanings of the
term Darwinism is to present a list of different interpretations of
the ter m , as encoun tered in the literature. In each case I will at­
tempt to analyze the validity of such usage and the temporal and
ideological context in which the term Darwinism was used with
this meanin g . This historical analysis will then permit us to ask
whether any one of the su ggested definitions of Darwinis m can
be singled out as the best one, or perhaps even the only cor­
rect one.

Darwinism as " Darwin's Theory of Evolution "

But which one is meant, since Darwin had so many theories of
evolution ? Should the term refer to the totality of Darwin's theo­
ries , including those of pangenes , the effect of use and disuse,
blendin g inheritance, and the frequency of sym patric speciation ?

. 92 .
 What ls Darwin ism?

Surel y not. To call such a conglomerate Darwinism would be

worse than useless ; it would be utterly misleading .

Darwinism as Evolutionism
Evolutionism was a concept alien to the ph ysicists , not only ow­
ing to its rej ection of essentialism but also for its acceptance of the
historical element, so conspicuously missing from the physics of
the mid-nineteenth centu ry. Historical inferences were equally
alien to all philosophers coming from logic or mathematics . It
was Darwin who made evolutionary thinking a respectable con­
cept of science. Nevertheless , it would be misleading to refer to
evolutionism as Darwinis n1 . Evolutionary thinking was already
widespread when Darwin published the Origin , particularly in
linguistics and in sociology (Toulmin 1 972 : 3 26) . For its existence
in biology one onl y needs to mention the names of Butron, La­
marck, Geoffroy, Chambers , and several German authors .
Clearly, Darwin was not the father of evolutionism, even though
he eventuall y b rought about its victory.

D a rwinism as Anticreationism
This is the Darwinism which denied the constancy of species and ,
in particular, special creation, that is , the separate creation of
every feature in the inanimate and living world . There were two
very different groups of anticreationists . The deists maintained a
belief in God b ut made him a rather remote lawgiver, who did
not interfere with any specific happening in this world, having
already arranged for everything through his laws . Whatever hap­
pened during evolution was the result of these laws . This thou ght
made evolution palatable to a number of Christian scientists such
as Charles Lyell and Asa Gray. However, only transformational
evolution-the orderly change in a lineage over time, directed
toward the goal of perfect adaptation-is susceptible to this deis­
tic interpretation. Darwin's variational evolution, with its ran-

. 93 .
 What Is Darwinism?

dom components at the level of both genetic recombination and

selection, cannot be insttuiuented by strict laws. The agnostic
anticreationists explained all evolutionary phenomena \Vithout
invoking any supernatural agents.
Immediately after 1 8 5 9 the word Darwinism simply meant a
rejection of special creation. If someone rejected special creation
and adopted instead the inconstancy of species, comn1on descent,
and the incorporation of man into the general evolutionary
stream, he was a Darwinian. Neither natural selection nor any
special theory of speciation, nor even one's belief in gradual ver­
sus saltational evolution, had any relevance to whether at that
time one was considered a Darwinian or not.
There were great differences between Darwin and other "Dar­
winians," such as Huxley, Lyell, Wallace, and Gray, on other as­
pects of evolutionary theory. But these differences were of minor
importance in the 1 86os, because the foren1ost meaning of Dar­
winism at that time was the rejection of special creation, together
with the adoption of the inconstancy of species, the theory of
common descent, and (excepting Wallace) the incorporation of
n1an into the animal kingdom. When someone in the 1 86os or
1 8 70s attacked Darwinism, he did so primarily in defense of cre­
ationisn1 or natural theology against these four Darwinian con­
cepts.
Perhaps the best way to determine what we should consider to
be the gist of Darwinisn1 would be to determine what Darwin
had in n1ind when he said the Origin was "one long argument."
Gillespie ( 1 9 79), in a careful analysis of this question, concluded
that it was Darwin's argument against special creation. Indepen­
dently of Gillespie I had come to the same conclusion when , in
the process of making a new index to the On�1?itz, I saw on how
n1any pages Darwin reiterated his conclusion that a particular
phenon1enon could not possibly be explained by special creation.
Instead Darwin argued for a n1aterialistic-that is, natural-ex­
planation for the diversity of the organic world and its history.
What Darwin pointed out again and again \Vas that any gi ven

· 94 .
 Wh at Is Darwi11 ism ?

phenon1enon for which s pecial crea tion had been invoked co uld
be explained n1 uch better by his theo ry of co n1 1non descen t .
Biogeogra phy was o f particular i1n portance i n that connection ,
because no other evidence is as convincing for co n1 n1on des cen t
as the distribution of species and higher taxa . For this reason l)ar­
win devoted two whole ch a pters of the Or(gi11 to biogeograph y�
and in these cha pters he sh owed over and over how a particular
distribution pa ttern could readily be explained by co n1n1on de­
scen t but not by special crea tion . It was the theory of co n1 n1on
des cen t which had the great unifying ca pacity about which l)ar­
win talked so o ften (Kitcher 1 9 8 5 : 1 7 1 , 1 8 4- 1 8 5 ) , because it at
once gave meaning to the Linnacan hierarchy, the a rchetypes of
the idealis tic m o rphologists , the history of biota, and many other
biological pheno men a .
C u riously, m any n1odern authors have claimed that Da rwin's
" one long a rgumen t " was an argun1ent in favor of natu ral selec­
tion (Recker 1 9 8 7) . There is no good evidence for this interpre­
tation . In his corres pondence Darwin referred to his manuscript
always as his " s pecies book ," not his book on natural selection .
Natural selection is dealt with in the first four chapters, appar­
ently in order to satisfy the vera ca usa demand of the leading phi­
losophers of science (Hodge 1 98 2) ; in the remaining ten chapters
natural selection is not featured . Instead, these chap ters are al­
most exclu sively devoted to docu mentations for common de­
scen t . I ndeed, D a rwin himself repeatedly called atten tion to " the
later cha pters " of the Origin as pa rticularly convincing proof of
his theory. The facts confi rm D arwin's claim . It was the reading
of these chapters that converted D arwin's contemporaries to a be­
lief in the incons tancy o f s pecies and the validity of com mon de­
scent, hence in evolution . Neither these cha pters nor the first four
chapters , however, produced many a dherents to his theory of
natural selection , a theory not adopted even by so1ne of Darwin's
closest frien ds and fo llowers , s uch as Hu xley and Lyell . This is
not so su rprisin g once one realizes that the Origin is not one long
argu ment in favor o f n atural selection .

· 95 ·
 ivhat Is Da rwin ism?

The adoption of evolution. ' by natural selection necessitated a

complete ideological uphe�val . The " hand of God" was replaced
by the wo rking of natural processes . God was "dethroned, " as
one of D arwin's critics formulated it. Indeed , God did not play
any role in Darwin's explanatory schemes .
B y what did D arwin replace him? What were the forces that
played the same role in D arwin's explanations that God played in
Christian dogma? Some physicalists , as well as historians and
philosophers adopting physicalist explanations , su ggested that
Darwin had adopted Newton's redu ctionist explanation - that
the processes of the inanimate as well as of the living world con­
sist of " a lawbound s ystem of matter in motion " (Greene 1 9 87) .
This wording fails to reflect Darwin's thinking . The biological
explanations adopted by Darwin are a long way fro n1 the New­
tonian explanation, and one looks in the Or(� in in vain for any
reference to " matter in motion ."

Da rwinis m as Anti-ideolog y
Not only natu ral selection but also many other as pects of Dar­
win's paradig m were in com plete opposition to n1 any of the dom­
inant ideologies of the mid-nineteen th century, as we have seen .
In addition to the belief in special crea tion and the design argu­
ment of natura l theology, other ideologies that were in total op­
position to D arwin's thinking were essentialism (typology) ,
physicalism (reductionism) , and finalism (teleology) . The adher­
ents of these creeds saw in Darwin's work their most formidable
opposition, and whatever the Orig in said or im plied that was dan­
gerous to their own position was designated by them as Darwin­
is n1 . But one by one these three ideologies were defeated , and
with their den1 ise the concepts of detern1inism, predictabili ty,
progress, and perfectability in the living world were weakened .
One of the by-products of the total refutation of all finalistic
aspects of o rganic evolution was the inevitable rein terpreta tion of
evolu tion as a historical process subj ect to ten1 porary con tingen­
cies . This led to the stress on opportunisrn in selection and on the
tin kering as pects of evolution . Such a concept of evolution is to-
 What Is Da rwin ism ?

tally d i fferen t fro n1 a l l s i n1 p l e tran s fo nnational chan ges in the in­

anin1 a te wo rld , wh ich , though also affected by cha nce, are p ri-
1n ari l y con t rolled by natural l a ws and thus permit ra ther st rict
p redi ction s . A closer a p p roach is provided by co mp lex inanin1ate
sys te1n s , s u ch a s the wea ther s ystem s , ocean cu rren ts (greatly af­
fected b y turbu lence) , and the interaction of continen tal plates
(res ulting in earthq uakes and vol canic eru ptions) , where the n1 ul­
titude of intera cti n g fa ctors and the frequency of stochastic pro­
cesses defy p rediction .

D arwinisn1 as Selectionism
A l m o s t any modern biolog i s t , when a s k e d what t h e term Dar­
winis m s tands for, will ans wer that it stands for · a belief in the
im portan ce of natura l selection in evolution. This interpretation
o f Darw inis m is so widely accepted today that it is someti mes
forgotten how rela t i vely new this modern version is . Natural se­
lection as the mechanism of evolutionary change was not univer­
sally a d o p ted by biologists un til the period o f the evolu tionary
synthesis ( 1 9 3 os-4os) ; however, natu ral selection al ways wa s the
key theory in D a rwin's total research program fo r at least some
evo l utionis ts . The firs t one of whom we know that this was true
was August Weis m ann (see Chapter 8) , but he was enthusiasti­
cally followed by A . R. Wal l a ce, who named his book on D ar­
winian evolution Da rwin ism because, as he said, " m y work tends
fo rceably to illus trate the overwhel ming i m portance o f natu ral
selection over all agen c ies in the p roduction of new s pecies . "
However, it t o o k another fifty yea rs a n d the refut ation of t h e ma-
j or anti-Da rwinian theories befo re this in sight was generally
adopted .

D arwinis m as Variational Evolu tion

D a rw in's con cept of evolu tion was ra dically different from the
transfo r m a tional and the saltational concepts of evolution that
had p revio u s l y b een p ro posed (see Chapter 4) . A lthough it

· 97 ·
 What Is Darwinism?

should have been obvious frqm the very beginning how different
Darwin's concept was, thi's· difference was not fully appreciated
until very modern times. Darwin's variational evolution was
based on entirely new philosophical concepts, and the neglect of
these concepts is the reason why it was usually confused with the
prevailing evolutionary ideas. Kitcher's discussions indicate how
far even some modern philosophers are from fully understanding
the magnitude of Darwin's departure from conventional think­
ing. Kitcher claims, " The trouble is that the theory I have as­
cribed to Darwin is uncontroversial-so uncontroversial as to
border on triviality" (1985 :3 0). Kitcher claims that virtually all of
Darwin's opponents accepted Darwin's statement that there is
variation among the n1en1bers of the species and that different
organisms have different properties. However, what Kitcher fails
to acknowledge is that the essentialist, as was en1phasized by
Lyell, Sedgwick, Herschel, and others, believes that there is a def­
inite limit to the a111ount of variation possible within a species or,
to put it differently, permissible to a single inherent essence. For
the population thinker, variation is unlimited. Hence, there is a
definite possibility of going beyond the confines of an existing
species. The nature and extent of variability was the crucial differ­
ence between the population thinker Darwin and his essentialistic
opponents. To call this difference uncontroversial completely ig­
nores its revolutionary importance.

D a rwinism as the Creed of the Da rwinians

Thwarted by the complexity of the Darvvinian paradign1 in their
atte1npts to reach a satisfactory definition of Darwinism, a nun1-
ber of historians and philosophers have attempted in recent years
to define Darwinism as the creed of the Darwinians. Hull (198 5)
and Kitcher (1985), in particular, have adopted this approach.
This choice of defining Darwinism is more often favored by phi­
losophers and historians than by biologists. It seems that a philos­
opher, particularly if he or she is coming from the background of
logic or mathematics, feels on safer ground discussing a sociolog­
ical phenomenon like the Darwinians than a conceptual frame-
 What Is Darwinism ?

work like l) arwini s n1 , which requires a thorough knowledge of

evolutionary biology. They attempt to j ustify thei r approach by
clai n1ing that Darwinism can best be dem arcated by the scien tific
con1 n1unity (the Darwinians) who su pport this systen1 . How­
ever, Recker ( 1 990) has pointed out how q ues tionable this cl ain1
is . Indeed , in s o n1c respects , this approach co mpounds the diffi­
culties because it is as difficult to define a Da rwin ian as it is to
define D arwinis 1n . For Hull, " the Darwinians fo rn1ed a fairly co­
hesive social g rou p . " Yet , this grou p p resum ably included only
Lyell, Huxley, and Hooker, because Asa Gray wa s in An1erica ,
Wallace was in the East Indies , Fritz Miiller was in South A mer­
ica, and Haeckel was in Germany.
Recker ( 1 990) , Hull ( 1 98 5 ) , and others have stated repeatedly
that there are no Darwinian tenets that cha racterize all the l)ar­
winians . I ndeed, it is true that some of the leading Darwinians,
like Huxley a n d Lyell , never believed in natural selection; neither
H u xley nor, p resu mably, Lyell en dorsed Darwin's complete
gradualis m ; neither Wallace nor Lyell thought that human beings
could be dealt with in the s a me way as animal species . Thus , there
were drastic differen ces among all of these Darwinians . This dis­
parity led H ull to state that it was not suffi cient for a person to
hold certain Darwinian ideas in order to be called a Darwinian . In
fact, he and others implied there was not a single concept sub­
scribed to by all of the so-called Darwinian s .
This i s a n error. There i s indeed one belief that a l l true original
Darwinians held in com mon , and that was their rej ection of cre­
ationis m , their rej ection of special creation . This was the flag
around which they assem b led and under which they ma rched .
When Hull claimed that " the Darwinians did not totally agree
with each other, even over essentials" ( 1 9 8 5 : 7 8 5 ) , he overlooked
one essential on which all these Darwinians agreed . Nothing was
more essential for them than to decide whether evolution is a nat­
ural pheno menon or somethin g con trolled by God . The convic­
tion that the diversity of the natu ral wo rld was the result of natu­
ral processes and not the work of God was the idea that brought
all the so-called Darwinians together in spite of their disagree­
ments on other of Da rwin's theories , and in spite of the retention

· 99 ·
 What Is Darwinism?

by some of them (Gray, Wallace) of other theological arg un1ents .

This situation was quite weJl' understood in the post-Origin pe­
riod and that is why at that time, for Darwin's opponents, Dar­
winism simply meant denying special creation and replacing it
with the theory of evolution and in particular the theory of con1-
mon descent.
The theory of evolution by natural means was powerfully sup­
ported by the explanatory power of the theory of con1 mon de­
scent. Indeed , it was this theory which eventually brou ght even
the morphological idealists into the Darwinian camp when they
realized that this was the only reasonable way to explain the hier­
archical arrangement of morphological archetypes . To be sure,
some morphologists, like Louis Agassiz , ascribed this order of
nature to God's laws. But the natural explanation of common
descent by D arwin and his followers was so much more convinc­
ing that Agassiz's interpretation fell on deaf ears and was no
longer heard after Agassiz's death in 1 8 73 .
The criteria by which one delimits scientific communities must
be ranked according to their importance. Creation or not was the
overwhelmingly important consideration in 1 8 5 9 . And it was the
adoption or rej ection of Darwin's thesis of evolution by natural
means that neatly separated the Darwinians from the non­
Darwinians. They did not need to form a closely knit social
group and might reside in Europe, South America, or the East
Indies, but they were held together and easily recognizable by a
single firm belief, that of evolution by natural means. And this
explains what has puzzled some historians, that so many
nineteenth-century evolutionists considered themselves to be
Darwinians even though they had adopted explanatory mecha­
nisn1s that were quite different from Darwin's natural selection.
Only the beliefs they shared with Darwin were considered by
then1 the truly crucial aspects of Darwinism .
To be sure, there were some borderline cases, like Asa Gray,
who seems to have adopted all of Darwin's theories and yet still
thought that God ultimately controlled everything , including the
nature of variation that was available to natural selection. Richard

• 1 00 •
 What Is Darwin ism ?

Owen was another borderline case because he really did believe

in evolution of some sort , but he thought he needed to attack
Darwin's theories uncompromisingly, in part owing to his en­
mity with Huxley.
Finally, there is the puzzling case of Lyell ( Recker 1 990) . Even
though a friend and mentor of Darwin and usually considered a
Darwinian, he never accepted the most basic components of Dar­
win's research program. God, for him, was apparently al ways the
ultimate causation; Lyell did not extend evolution to man, and he
never accepted natural selection. If one wants to explain why
Lyell continued to support Darwin even though he disagreed
with mos t of Darwin's evolutionary beliefs , one must not forget
that Darwin originally was a geologist-not only a geologist but
very definitely a Lyellian geologis t, who had strongly supported
Lyell in his geological writings . It is quite possible that Lyell
simply reciprocated, through his support of Darwin's ideas , for
everything Darwin had done in his geological writings to support
the Lyellian view. And, as Hodge has pointed out so convinc­
ingly, Da rwin was a Lyellian when he began to occupy himself
with biological problems , even though ultimately he rebelled
against some of the most basic beliefs of his teacher. Thus , polit­
ically and social ly, Lyell belonged to the Darwin party. Concep­
tually, he never was a real Darwinian.
Except for these borderline cases , the issue was absolutely
clear-cut for Darwin's contemporaries . If someone believed that
the origin of the diversity of life was due to natural causes , then
he was a Darwinian. But if he believed that the living world was
the product of creation, then he was an anti-Da rwinian. The ex­
istence of a few deistic borderline cases is no more a refu tation of
this basic classification than the exis tence of a number of incipient
species is an argument agains t the existence of species .

Darwinis m as a New Worldview

J.C. Greene ( 1 9 8 6) has sug ges ted, in line with the view of other
historians of ideas , that the affix - ism should be used only for ide-

• 101 •
 What Is Darwin ism ?

ologies and not fo r scientific theories . I agree with him that one
.'
should not dignify adheret1ce to ordinary scientific theories with
the ending -ism . However, there are scientific theories that have
beco me im portant pillars of ideologies , as is the case in Newton­
ianis m , and this is certainly true for Darwinis m . Some of Dar­
win's important new concepts , like variational evolution, natural
selection , the interplay of chance and necessity, the absen ce of
supernatu ral agents in evolution, the position of man in the realn1
of life, and others , are not only scientifi c theories but are at the
same time important philosophical concepts , and characterize
worldviews that have incorporated these concepts . Thus , as far as
several of Da rwin's most basic scientific theories are concerned ,
they have a legitin1ate standing both in science and in philosophy.
The rej ection of special creation signified the destruction of a
p reviously ruling worldview. This is why not onl y scientists like
Sedgwick and Agassiz, bu t also philosophers like Whewell and
Herschel , opposed Da rwin so vigorously. Was there a new
worldview that took the place of creationis m ? If so, what was it
and how could it be defined ? Greene suggests that henceforth
" the word Darwinism should be used to designate a world view
that seems to have been arrived at more or less independently by
Spencer, Darwin , Huxley, and Wallace, in the late I 8 5 0s and early
1 860s ." Here he refers to a Victorian worldview in which certain
sociological ideas were used to develop a new social theory. It was
based in part on the writings of Adam Smith, Malthus , and Da­
vid Ricardo, and postulated that co mpetition , struggle, and the
increase in populations would result in progres s . Darwin was fa­
n1iliar with these ideas, but as has been shown by various ca reful
analyses of the writings of Darwin and the social philosophers ,
these idea s were not the sou rce of Darwin's biological ideas (see
Gordon 1 9 89) , as much as some political writers \vould want us
to believe this .
In his writings Darwin never upheld such a wo rldvie\v. Being
unable to find any support fo r his clain1s in Darwin's \v ritings ,
Greene suggests that we adopt Herbert Spencer's \vorldvie\v,
which, according to Greene , was very much the san1e as that of

• 1 02 ·
 Wha t Is Darwinism ?

D a rwin . S pencer's worldview is described by Greene as follows :

"As a law bound s yste n1 of matter in motion , evol ution ary deism
verging towa rd a gnosticis111 under the influence of posi tivistic
empiri ci s m , the idea of organic evolu tion , the idea of a social sci­
ence o f histori cal developn1 ent, faith in the benefi cient effects of
co m petitive s t ru ggle," to single out the n1ost im portant points .
Was this really Darwin's worldview?
There a re indications that D arwin shared in the prevailing be­
liefs o f many enligh tened u pperclass Englishmen , includin g (ac­
cording to Greene) S pencer, H uxley, and Wallace. But looking
critically at the lis t of these beliefs , as presented by Greene, one
dis covers that it does not include a single one that was original
with D arwin , in fact, not even a single one original with S pen cer.
Indeed , most o f these views go all the way back to the eigh teenth
century, even thou gh some of them had changed their meaning ,
such as the term " st ruggle for existen ce," which already had been
part o f natu ral theology. E ven thou gh Greene adopts the term
" Da rwinism " for this set o f ideas , he adn1its , " S pencer could
rightfully demand that it be called Spencerianism , " because it
does not include a single one of Darwin's own ideas. What is
worse, this S p encerian paradigm is in several respects in com plete
conflict with D a rwin's idea s . Fo r instance, S pencer supported
transformational rather than variational evolution ; second, his
evolution was dis tinctly teleological ; and finally, it was based en­
tirely on an inheritance of a cquired chara cters , not involving nat­
ura l selection in any manner. Hence, it was not only scientifically,
but also philosophically, something quite different from Darwin's
set of ideas. To claim that Darwin and Spencer supported the
same paradigm is a clear falsification of histo ry. It is a popular
thesis with sociologists , but biologists who have looked into this
problem in recent years have been unanimous in refu ting it (Free­
man 1 974) .
How lon g did the " D arwinis m , " as defined by Greene, prevail
as a world view ? According to Greene, through the 1 86os , and
perhaps u p to about 1 8 7 5 , but Spencer, Wallace, and Huxley ac­
tually abandoned one plank or the other of this platform soon

. 1 03 .
 Wh a t Is D arw in ism ?

after the 1 86os . The conseq l}�nces of scien tific Darwinism made
the acceptance of this soci·a1 theory of Darwinism quite un ten­
able.
Greene su ggests that certain modern biologis ts , like Julian
Huxley, George Gaylord Simpson , and perhaps Edward 0 . Wil­
son , have an updated Darwinian worldview. The truth of the
matter is that unless a person is still an adheren t of creationis n1
and believes in the literal truth of every word in the Bible, every
modern thinker-any modern person who has a wo rldview -is
in the last analysis a Darwinian . The rejection of special creation ,
the inclusion of man into the realm of the living wo rld (the elim­
ination of the special position of man versus the anin1als) , and
various other beliefs of every enlightened n1odcrn person are ul­
timately all based on the consequences of the theories con tained
in the Origin o_f Species . Nevertheless, to define Darwinism as the
world view sup posedly held by Darwin in the I �oos would be
about the most useless definition I can in1 agine .

Darwinis n1 as a New Methodolo g y

In view o f the intense preoccupation of modern philosophy of
science with questions of methodology, it is not surprising that
several philosophers have asked themselves what Darwin's
method of science was , and what was new in it. Was Darwin's
1nethod strictly hypothetico-deductive , as sug ges ted by Ghiselin
( 1 969) , or did he follow various other schemes ? There have been
diverging answers to these questions (Recker 1 9 8 7) . Almost any
modern philosopher of science has su gges ted a somewhat differ­
ent Da rwinian n1ethodology. How inductive was Darwin ? Does
the se1n antic approach to the philosophy of science describe Dar­
win's ap proach best? These are the sorts of questions being asked .
Da r\vin realized that to convince his readers of the validity of
his concepts he had to adopt a n1ethodology that was rather dif­
ferent fro n1 that used by physicists to demonstrate the validity of
universal p h ysical laws . Darwin's method was to present the evi­
den ce on which he based his inferences , and he used these infer-
 Wlz at Is Da rwin isn1 ?

ences to support his conjectures. The greater the nu1nbcr and the
variety of pieces of evidence he could cite, the n1ore convincing
the inferences bccan1 c (Ghisclin 1969). Kitcher ( 19 8 5) belittled
Darwin's 1nassive citing of supporting facts, not understanding
that these facts were of little interest as such but only as evidence
and docun1entation for the inferences that Darwin was 1naking.
These inferences not only served to support his conjectures, but
they also reveal son1e of l)arwin's basic underlying ideas.
One of the reasons why different authors were able to claim
that Darwin had used different 1nethodologies is that l)arwin was
rather pluralistic, n1ethodologically. In son1e argu1nents he did
indeed follow the hypothetico-deductive 1nethod; in others he
was proceeding in an inductive 1nanner. I think that any claim
that Darwin consistently applied only a single method could eas­
ily be refuted. And what is more in1portant, the ultimate valida­
tion of most of Darwin's theories did not result from the victory
of his n1ethodology but fron1 additional facts and the gradual ref­
utation of opposing ideologies.
Many authors have called attention to the spectacular unifying
capacity of Darwin's paradigm. As Dobzhansky stated it, " Noth­
ing in biology n1akes sense except in the light of evolution" ; I
might modify this by saying "in the light of Darwinian evolu­
tion. " I think it is quite misleading to suggest, as is done by
Kitcher, that it was the goodness of his methodology that had this
unifying effect.
There is a widespread belief among philosophers that a theory
has virtually no chance of being accepted unless an appropriate
mechanism is proposed sin1ultaneously. This is indeed often true.
Wegener's theory of continental drift was not accepted by the
geophysicists until the mechanism for the movement of conti­
nents had been elucidated in the theory of plate tectonics. It is not
necessa ril y true, however, as shown by the theory of common de­
scent. Darwin's proposed mechanism, natural selection, was al­
most universally rejected, but since the fact of evolution and the
theory of common descent were so completely convincing after
Darwin had pointed them out, other evolutionists simply

• 1 05 •
 What Is Darw in ism ?

adopted-instead of naturaL selection-various other kinds of

mechanisms, whether teleological , Lamarckian, or saltational .
Indeed, for Darwin himself, as much as h e believed in natural
selection all his life, it was obviously not this mechanism that was
of first importance for him but the evidence for evolution and
common descent. Hence the completely unbalanced assignment
of space to these two subjects in the Origin .
The version of Darwinism that developed during the evolu­
tionary synthesis was characterized by its balanced emphasis both
on natural selection and on stochastic processes; by its belief that
neither evolution as a whole, nor natural selection in particular
cases, is deterministic but rather that both are probabilistic pro­
cesses; by its emphasis that the origin of diversity is as important
a component of evolution as is adaptation; and by its realization
that selection for reprodu ctive success is as important a process in
evolution as selection for survival qualities.

The Pluralis m of Darwinism

It is now clear that no simple answer can be given to the question
"What is Darwinism? " When someone asks this question, he is
bound to receive a different answer depending on the time that
has passed since 1 8 59 and on the ideology of the person that was
asked. Such pluralism is not congenial to many philosophers, and
they have been trying to find some method by which they could
attach the term Darwinism to a very definite meaning . The su g­
gestion was made , for instance, that one should select an " exem­
plar" to fix the meaning of the term Darwinism , analogous to the
action of a taxonomist who selects a type specimen in order to
anchor a species to a definite name . Hull ( 1 9 8 3 ; 1 9 8 5 ) has indeed
attempted to do so, but I have shown elsewhere ( 1 98 3 , 1 989)
what insurmountable difficulties oppose the application of the
exemplar method.
The majority of the nine meanings of the term Dar\vinism dis­
cussed above are clearly either misleading or unrepresentative of
Darwin's thought. Looking at the situ ation as a historian, I am

• 1 06 •
 What Is Darwinism ?

impressed that two meanings have had the widest acceptance.

After I 8 5 9, that is, during the first Darwinian revolution, l)ar­
winism for almost everybody meant explaining the living world
by natural processes. As we will see, during and after the evolu­
tionary synthesis the term "Darwinism" unanin1ousl y meant
adaptive evolutionary change under the influence of natural selec­
tion, and variational instead of transformational evolution. These
are the only two truly meaningful concepts of Darwinisn1, the
one ruling in the nineteenth century (and up to about 1 9 3 0), the
other ruling in the twentieth century (a consensus having been
reached during the evolutionary synthesis). Any other use of the
term Darwinism by a modern author is bound to be misleading.

• 1 07 •
 C H A P. J E R E I G H T
-------------• -------------

A H ard Look at Soft Inherit ance:

Neo-D arwinism

D AR w I N H I M s E L F was in part responsible fo r the diffi culties

his theories encountered in the years follow ing the publi­
cation of t he Or({!it1 . On most any subj ect he dea lt with-and this
incl udes nearly all of his theories-he not infrequently re versed
himsel f. One of Darwin's twen tie th-cen tury criti cs wa s not en­
tirel y unj ust in ass erting that " l)arwin's hedging and sel f con tra­
diction en able any unscrupulous reader to choose his text fro n1
the On�'< in o.f Sp ecies or Th e Desce11 t o.f A1a11 with aln1ost the san1e
ease or acco111 moda tion to his p u rpose as i f he had chosen fro n1
the Bible" ( Barzun 1 9 5 8 : 7 5 ) . For ins tance, although in principle
Da rwin rej ected essen tial isn1 and explained adaptation in strictly
variational terms (the individual was fo r hin1 the target of selec­
tion) , he son1eti mes fell back into typological langu a ge, as fo r the
origin of an inci pient s pecies by the isola tion of a variety. Da rwin
stron gly rej ected any teleological " law of neces sary develop­
men t " ( I 8 5 9 : 3 5 I ) , and yet he made a suffi cien t number of ca reless
sta te111ents to permit certain historians to con sider Dar\vin a te­
leol ogist ( H i m melfarb 1 9 5 9) . A l though he often stated that
enough variability was available fo r natural selection to explain
the origin of every adaptation, he nevertheless decl ared his puz­
z le1nen t over the evolution of eyes . On one page he n1 ight say
that only s light variations are evolutionarily i 1n portan t , but then
on another page he will discuss rather strikin gly different va rie­
ties ., like the an con sheep and the turns pit dog , both of then1 ex­
tre1nely sho rt-l egged . Kohn ( 1 989) has n1 ade a detailed anal ysis
of Dar\vin's vacillation and its psy ch olo gical and tactical rea­
so ns .

• 1 08 •
 A Hard Look at S(ft /11/zeritm1ce

It is frequently stated that Darwin totally rejected all Lan1arck­

ian ideas in the first edition of the Or({!in and that he allowed at
that tin1e no n1echanisn1s of evolution other than random varia­
tion and natural selection. This is not correct. l)arwin displayed
considerable indecision already in 1 8 5 9 on the origin of variation
and the nature of inheritance. He makes no less than three sets of
concessions to the possibility that the environment in the widest
sense of the word can induce genetic variation and that these ac­
quired characters can be inherited. First, he speculated about a
direct effect of the environn1ent on certain structures; second, he
hypothesized an indirect effect of the environment in increasing
variability ; and third, he discussed the effects of use and disuse,
for instance, when he says that the reduced size of the eyes of
moles and other burrowing man1mals is "probably due to grad­
ual reduction from disuse but aided perhaps by natural selection"
( I 859 : I 3 7). In the case of cave ani1nals, when speaking of the loss
of eyes, he says, " I attribute their ioss wholly to disuse." On the
other hand, he calls attention to the fact that the adaptations of
the neutral castes of social insects are inexplicable with any kind
of Lamarckian theory.
Darwin's maj or thesis was that evolutionary change is due to
the production of variation in a population and the survival and
reproductive success ( "selection") of some of these variants. But
the origin of this variation puzzled him all his life. Darwin con­
sidered variation to be an intern1ittent phenomenon, one that oc­
curred mostly under special circumstances. However, he was
quite sure that in nature there was an immense reservoir of varia­
tion that was always available as the material of selection. Darwin
devoted the entire fifth chapter of the Origin to the "laws of vari­
ation" and followed it up some years later with the two-volume
work The Variation of An ima ls and Plan ts under Domestica tion
(1 86 8). Two years earlier, unknown to l)arwin, Gregor Mendel
had published the key to an understanding of the nature of varia­
tion. But Darwin never heard of Mendel's work and was never
able to solve the problem. Un der the influence of the prevailing
concepts of physicalism, Darwin looked for a direct cause of each
variant. Also, his partial belief in blending inheritance-the fu-

• 1 09 .
 A Hard Look a t Soft In herita n ce

sion of the inheritable contribution

."
of the parents in the off-
spring-made it impossible· for Darwin to solve the problem. His
partly typological, partly deterministic approach precluded his
finding the right answers.
The great number of misconceptions about the nature of inher­
itance and the absence of good empirical data led one of Darwin's
younger contemporaries to conclude that the study of variation
was the new frontier of evolutionary biology. This young evolu­
tionist was August Weismann ( 1 8 3 4-1 9 1 4) . By his uncompro­
mising refutation of an inheritance of acquired cha racters, and by
his genetical theorizing , even when wrong , he laid the foundation
for the acceptance of Mendelian inheritance, the nex t major ad­
vance toward modern evolutionism.
In the first twenty years after the publication of the Origin no
conceptual contributions were made to the Da rwinian paradigm.
That is, none were having any impact . To be sure, Mendel pub­
lished in 1 866 his fundamental discoveries about inheritance, but
they were completely ignored (until 1 900) and left contemporary
evolutionary thinking unaffected. A decisive new departure did
not happen until 1 8 8 3 , when Weismann published his provocative
essay On Heredity ( 1 8 8 3 ) . Here, Weismann denied categorically
any occurrence of the effects of use or disuse, indeed any inheri­
tance of acquired characters. In this uncompromising position he
was immediately joined by A. R. Wallace, who pleaded for the
monopoly of selection in his book Darwinism ( 1 8 8 9) . George J.
Romanes, a disciple in Darwin's later yea rs, considered the denial
of soft inheritance a major departure from Darwin's own teaching
and therefore in 1 8 96 coined the term "neo-Darwinism" for
Weismann's exclusive selectionism, or, as Weismann himself once
pu t it, Darwinism without inheritance of acquired characters.
This term continued to be used throu gh the evolu tionary synthe­
sis, but once a belief in soft inheritance had been universally ab an­
doned , most people reverted to the simple term "Darwinisn1 ." In
the years between 1 8 8 3 and the evolu tionary synthesis, the fight
against soft inheritance was intin1ately connected \Vith the name
Weisn1ann.

• 1 10 •
 A Hard Look at Soft lnh erita)1ce

August Weismann is one of the towering figures in the history

of evolutionary biology. If we ask who in the nineteenth century
after Darwin had the greatest impact on evolutionary theory, the
unequivocal answer must be Weismann . What makes Weisn1ann's
concern with the problen1s of evolution so fascinating is the grad­
ual n1aturation of his thinking . There was an unfortunate ten­
dency in the older histories of science to depict the thought of a
great scientist as a n1onolithic structure, something poured out of
one mould, impressively unchanging. More recent researches
have shown how misleading such a presentation is, and this is true
for almost any scientist one may 1nention. W hen Weis1nann re-
jected the theory of an inheritance of acquired characters, surely
a dramatic conversion, he was already over forty-seven years of
age. This change required a fundan1ental rethinking of all his pre­
vious assumptions on inheritance and evolution.
I t may be helpful to recognize three periods in Weismann's
thinking : 1 86 8 - 1 8 8 1 or 1 8 8 2, during which he accepted the in­
heritance of acquired characters; 1 8 82- 1 895, when he searched
for a source of genetic variation; and 1 896 - 19 1 0, when he recog­
nized germinal selection as the aid of natural selection.
In 1 8 5 9 , when Darwin published his revolutionary On the Ori­
gin of Species, Weismann was twenty-five years old. The subject
of evolution was never raised during his student days at several
German universities. Nor did Weismann's early zoological work
have anything to do with evolution; it dealt instead with such
topics as muscle histology and insect embryology. And yet, in
1 86 8, Weismann chose Darwinism as the subject of his inaugural
lecture at Freiburg. In no other country, not even in England, did
Darwinism have the impact it had in Germany, as is evident from
the writings of contemporary zoologists, botanists, and anato­
mists. The high importance Weismann attributed to Darwin's
thought is documented by the fact that he compared the "trans­
mutation" theory to the Copernican heliocentric theory and im­
plied that no advance in human understanding since the accept­
ance of that theory had had as great an impact as Darwin's theory
( 1 86 8 : 3 0).

• 1 1 1 •
 A Hard Look a t Soft In herit ance

Weis m an n unders tood frow the beginning that one must dis­
tinguish t wo theories of evolution ; evolution as such , called b y
Weis m ann the t ransmutation theory; and Da rwin's explanatory
theo ry, the theory o f natural selection .
Weisman n's attitude toward evolution as such was close to that
o f the modern evolutionis t , for whom evolution is not a theory
but an accepted fact . The various con clusions arrived at by evo­
lutionary biology, s aid Weismann, " m ay be maintained with the
same degree of certaint y as that with which astrono m y as serts
that the earth moves a round the sun; fo r a conclu sion 1n ay be
a rri ved a t a s safely b y other methods as b y mathen1atical calcula­
tion " ( 1 8 8 6 : 2 5 5 ) . The fa ct of evolu tion was so irrefutably estab­
lished fo r Weismann that in h is subsequen t writings he virtuall y
never bothered to list fa cts in support of evolu tion but instead
concen trated on the causal as pects of the evol utionary proces s .
Weis mann en thus ias tically accepted the theo ry o f co m mon de­
s cent even though the constru ction o f phylogenies was not one of
his fields o f s pecializa tion . I n pa rti cular he fully en dorsed the
theory of reca pitulation and based his analysis of the ontogenetic
s tages of s ph in gid caterpillars en tirely on this prin ciple . Weis­
m ann wa s aware of the relative independence o f evol utionary de­
velop men ts in the on to gen y of different orders of in sects , b y call­
ing attention to the overall uniformity in the morphology of
adult was ps and flies and by interpreting the numerous s peciali­
zations found in the la rval sta ges as secondary adap tation s . ( For a
further discussion of Weisn1ann's views on recapitula tion see
Gould 1 97 7 : 1 02- 1 09 . )

Weis n1ann's Assault on Antiselectionis m

By fa r the n1ost i m po rtant of Darw in's evolutionary theo ries \Vas
natural selection . It a ttem pted to explain \Vith the help of ma terial
causes \vh a t had been explained previou sly by a supernatural
cause, by design . This theory \Vas so novel , so daring , that at first
it \Vas adopted by very fe\v biologists . Weisn1 ann \Vas one of the
few � and, as we shall see, even tually he \Ven t even fu rther than

• 1 12 •
 A Hard Look at S o_ft Inheritance

Darwin in asserting the " allsufficiency " of natural selection . Most

evolutionists in the post-D arwinian period supported other theo­
ries . The th ree n1ost widely accepted of these opposing theories
(Bowler 1 9 8 3 ) were ( 1 ) a belief in an intrinsic driving fo rce , or
" phyletic fo rce ," resulting in evolution by "orthogenesis " ; (2) sal­
tatio11al evolution ; (3 ) La1narckian factors (inheritance of acquired
cha racters ; sec bel ow) .
I n the 1 86os and 1 8 70s the idea of some teleological , finalistic,
phyletic force was perhaps the most popular of the three options ,
at least in Germ any. I n view of the fact th at it had been adopted
by some of Germany's n1 ost admired biologists, such as von
Baer, Naegeli, and Kolliker, b y the orthogenisists Haacke and Ei­
mer, and by leading philo sophers , particularly von Hartn1 ann ,
Weis1nann found it necessary to en1phasize again and again his
resistance to the accep tance of such a metaphysical force , and he
attem pted to refu te it by ever-new arguments . In view of the
prestige of his opponents , he chose not to ridicule this school of
thought but rather to show that in his opinion it was inconsis ten t
with the known facts . " How could such intrinsic drives produce
such sophisticated p atterns as found in leaf imitating butterflies ,
like K alli m a ? " he asked . Fu rthermore, if there were such an evo­
lutionary force, he said, it should be possible to establish it by
empirical research . But no one had succeeded in doing so; in­
stead , he sai d , it was possible to trace all evolu tionary changes " to
know n transforming factors ," that is, to natural selection
( 1 8 82 : 1 6 1 ) .
The second of the great antiselectionist theories of the post­
Darwinian period was the origin of new types and organs by ma-
j o r saltations . This contention was dia metrically opposed to evo­
lutionary gradualism , one of Darwin's fi ve evolutionary theories .
Yet saltationis m was favo red by T. H . Huxley, Kolliker, and other
Darwin con temporaries . Weis m ann was unalterably opposed to
the possibility of saltational evolutionary change ( 1 8 8 2 : 697) . It is
interesting that in 1 8 82 and 1 8 86 Weis mann already anticipated
the clai ms made a few years later by Bateson ( 1 8 94) and de Vries
( 1 90 1 ) and attem pted to refute them . "An abrupt transformation

• 1 13 .
 A Hard Look at Soft Inheritance

of a species is inconceivable,� pecause it would render the species

incapable of existence " ( 1 8.S'6 : 264) , he wrote, believing that the
existence of numerous coadaptations would make such an instan­
taneous total restructuring of the organis m an impossibility.
A fter all, when studying organis ms one is amazed at the precision
and ubiquity of adaptations . " It is evident that one can not possi­
bly explain these innumerable adaptations by the occu rrence of a
rare, accidental l y at one ti me occurring, variation. The necessary
variations with the help of which selection pieces together evo­
lutionary change must always be offered again and again in many
individual s " ( 1 892 : 5 68 ) . At the height of the popularity of de
Vries 's mutation theory, Weismann once more demonstrated the
utter improbability of saltational evolution ( 1 909 :22-24) .
Weismann was prou d to be able to assert, " more definitely than
Darwin has done," that all changes must have occu rred " very
gradually and by t_he smallest steps " ( I 8 8 6 : 264) . It is obvious why
Weis mann had to insist on gradual evolutionary change when one
remem bers that at first he attributed much of it to use and disuse,
and later considered evolutionary change as more or less due to
quantitative changes in large num bers of determinants . Even
after de Vries had published his mutation theory, Weismann pos­
tulated that a saltation of the phenotype was simply the making
visible of a long series of preceding small genetic changes ( 1 904,
2 : 1 1 9) .

Weis mann's Evidence for Selection

Rej ecting all other possible causations for evolutionary change,
Weismann firmly adopted selectionis m . He showed himself a
convinced selectionist in his very first evolutionary publication
( 1 8 68) . He supported his selection ism by various sets of argu­
ments . In view of the evident invalidity of all teleological and
saltational theories of evolution , said Weismann , there is really no
other option than to accept natural selection . There is not a detail
in the structure or physiology of an organism tha t has not been

• 1 14 .
 A Hard Look at Soft In heritance

shaped b y natural selection . Without a doubt, Weismann was the

most consis tent selectionist in the nineteen th century.
Natu ra l selection was the main topic of the series of papers
Weis m ann published in the 1 8 70s and 1 8 8os dealing with the
theory o f descen t . In his detailed studies of the markings of
sphingid caterpill a rs, he unequivocally adopted the adaptationist
progra m . He asked : " Have the n1arkings of the caterpillars any
biological value, or a re they in a measu re only sports of natu re?
Can they be considered as partially or en tirely the result of natural
selection ? Or has this agency had no share in their production ?"
( 1 8 8 2 : 3 08 ) . To be able to answer these questions , Weism ann de­
veloped a rigorous methodology. And he tes ted his assu mptions
with carefully execu ted experiments in which caterpillars that
had been placed on a variety of backgrounds were exposed to
birds or liza rds . Furthermore, Weisn1ann always sought to deter­
mine whether or not there was a correlation between structural
or color characters and behavioral characteristics . The result of
these s tudies was that he found abundant evidence in favor of se­
lection . " I t has been possible to show that each of the three chief
elements in the markings of the S phingidae have a biological sig­
nificance, and their origin by means of natural selection has thus
been made to a p pear pro bable" ( 1 8 82 : 3 80) .
Wherever Weis m ann looked , he found evidence for selection­
not only in the animal kingdom but also among plants . He ex­
amined the adapti ve significance of the form and color of flowers ,
so beautifully demonstrated by D arwin for orchids and by nu­
merous o ther s tudents of flower biology beginning with S p ren­
gel ; the ven ation of leaves ; and the numerous protective devices
of plants against herbivores . Weismann p resented a su perb de­
scripti on of the aquatic adaptations of marine mammals, demon­
stra tin g that all the differences between them and terrestrial m a n1-
mals a re clearly adaptations ( 1 8 86 : 26 1 ) .
Natu ral selection is active not only in the acquisition of new
adap tations but in the maintenance of existing ones . As soon as
the selection p ressure is relaxed, as in the case of the eyes of cave

• 1 15 .
 A Hard Look at Soft Inheritance

animals, individuals with imperfect structures will no longer be

eliminated. " In my opinion every organ is kept at the peak of its
conformation only by continuous selection. And it slides down
from this height ceaselessly even though very slowly, as soon as
it no longer is of value for the survival of the species'' ( 1 893 : 5 1 ).
Such a loss of organs can result from either a relaxation of the
maintenance selection or from an actual counter-selection exer­
cised by competition for tissue substrate. Weisn1ann presented a
particularly impressive documentation of natural selection in his
Vortrage ( I 904 I ; 3 6 - 1 70).
Weismann professed without hesitation that he was a panselec­
tionist. " There is no part of the body of an individual or of any of
its ancestors, not even the minutest and most insignificant part ,
which has arisen in any other way than under the influence of the
conditions of life" ( 1 8 86 : 260). He admitted, however, that " these
are indeed only convictions, not real proofs. " In the 1 88os direct
experi1nental proof for the force of selection was scanty. The
analogy with artificial selection was still perhaps the most con­
vincing evidence for selection; for, as Weismann said, the breed­
ers had achieved almost any objective of their selection ( 1 89 3 : 60).
Otherwise the principle of exclusion is the best support for natu­
ral selection, a principle which states that a theory can be adopted
for the time being if all competing theories have been refuted.
After Weisn1ann had become an ardent selectionist, he realized
(as had Darwin before him) that selection does not necessarily
lead to perfection. " No device in nature is absolutely perfect, not
even that beautifully constructed eye of man. Everything is only
as perfect as necessary, at least as perfect as it must be in order to
acco1nplish what it is supposed to accon1plish. '" This echoes si1n­
ilar state1nents n1ade by Dar\vin ( 1 859 :20 1 , 206).
In recent years several authors have proudly announced the dis­
covery of constraints on the action of natural selection. Weis1nann
was aware of the role of developn1ental constraints more than a
hundred years ago, a sign of his deep understanding of the evo­
lutionary process. The physical constitution of each species lin1its
the action of natural selection because it urestrains the course of

• 1 1 6 .
 A Hard Look at S,ft Inheritance

develo p n1ent, however wide the la tter may be" ( 1 8 8 2 : 1 1 3 -1 1 4) .

These develo p 1n ental constraints arc the reason " why organic
evolution has frequently proceeded for longer or shorter periods
along certain develop men tal lines " ( 1 8 86 : 2 5 8) .
The ques tion of what the target of selection is occu pied Weis-
1nann all his life . At first he followed Darwin , recognizing only a
sin gle level o f selection, and evidently consid ered the individual
as a whole to be the ta rget . He considered the genotype as a holis­
tic syste m , the individual co n1 ponents of which could not be ran­
do m l y interchanged or replaced . To what extent Weisn1 ann gave
up his holistic view after he had developed his co1n plex theories
of the structure of the genotype, and his postulate of three levels
of selection, has not yet been determined .
As for sexual selection , Weismann at first was quite enthusias­
tic, and in 1 8 72 he attributed many differences between the sexes ,
even in i nsects , to sexual selection ( 1 8 7 2 : 60) ; indeed he felt the
theory should be applied even more extensively. Weis mann saw
clearly that sexual selection gave an advantage to certain individ­
ual s , and he rem a rked that the driving force of sexual selection is
not the external environ ment but rather the preferences of the
individuals in their selection of a mate . Characters acquired by
sexual selection do not o ffer any advantage in the daily struggle
for exis tence ( 1 8 72 : 62) .
Ten years later Weismann ( 1 8 8 2 : 1 0 1- 1 02) had become rather
con fused about the cause of sexual dimorphis m . Twenty years
later Weism ann returned co mpletely to Darwin's position . He de­
voted an en tire chapter of his Vortraj?e ( 1 904 chap . 2) to sexual
selection , definitel y accepting the p rinciple of female choice and
attem pting to refu te all o f Wallace 's obj ections ..
The su bsequent history of evalu ation of sexual selection has
been one o f ups and downs . When the mathematical population
geneticists decla red the individual gene to be the unit of selection
and defined fitness as the contribution of such genes to the gene
pool of the next generation , there was little room left for sexual
selection . This attitude characterized the perio d from the 1 920s to
the 1 960s . In the past two decades the importance of sexu al selec-

• 1 1 7 .
 A Hard Look at Soft Inheritance

tion has again been acknowledged; it has been recognized that the
individual as a whole is the principal target of selection and that
such an individual might have a reproductive advantage owing to
characteristics that do not contribute to general fitness. This topic
has become one of the major concerns of sociobiology.

Rej ection of the Inheritance o f Acquired Characters

When Weismann concluded in 1 8 8 2 that the theory of an inheri­
tance of acquired characters was untenable, he had to search for a
new source of genetic variation and therefore devoted the second
research period mostly to genetic studies. Indeed, he stated quite
explicitly that his theory of inheritance "was so to speak only a
means toward a higher purpose, [ to establish] a foundation for
the understanding of the transformation of organisms in the
course of time" ( 1 904:iii). Weismann understood from the very
beginning that variation was an indispensable prerequisite for the
operation of natural selection. Variation was no problem for him,
because at that time he believed in "the origination of transfor­
mations by the direct action of external conditions of life"
( 1 8 8 2 :6 8 2). There is probably no other evolutionist whom histor­
ians traditionally have considered as extreme a selectionist as
Weismann. So it will come as a great surprise when they learn
that, at least until 1 8 8 1 , Weismann, like Darwin, believed in an
inheritance of acquired characters. Not only did he believe in the
heritable effect of use and disuse, but he stated as late as Novem­
ber 1 8 8 1 ( 1 8 8 2 :xvii) : "Nor can the transforming influence of di­
rect action, as upheld by Lamarck, be called in question, although
its extent can not as yet be estimated with any certainty." (Weis­
mann's conversion to "hard" inheritance must have occurred be­
tween 1 8 8 1 and 1 8 8 3 . He still supported an inheritance of ac­
quired characters in at least three statements in the preface , dated
November 1 8 8 1 , to the English edition of his Studie11 zu r Desce11-
de11ztlz eorie, 1 8 76, published in I 8 8 2. In this edition Weismann
added a number of footnotes on various subj ects but rnade no
disclaimers to some of the strongly Lamarckian statements of the

• 1 1 8 .
 A Ha rd Look at S(ft Inh eritance

original Gennan text. ) That variation was an indispensable com­

ponent of the process of natural selection ren1ained Weismann's
firm conviction even after he had changed his mind completely
about the causation of such variation. Even in 1 893 , he still re­
ferred to variation "as one of the main factors of natural selec­
tion" ( 1 893 :54).
The inheritance of acquired characters was for Weisn1ann the
major source of variation in all of his evolutionary writings prior
to 1 882 . He later ad1nitted, "Twenty-five years ago I was still of
the opinion that in addition to [ other factors] also the inherited
effect of use and disuse played a not unimportant role in evolu­
tion" ( 1 893 :43 ). Actually, in Weismann's pre- I 8 83 writings one
can find numerous suggestions about how the environment
might influence the variation and inheritance of organisms
(Churchill 1 96 8; Blacher 1 9 82). What Weismann envisioned was
that species and populations went through occasional periods of
greatly increased variability, and that if segments of a population
were isolated during such a period, then a process he called
"amixia" would lead to differences. Some of Weismann's descrip­
tions sound like what later was called "genetic drift."
Because this is often misunderstood, let me emphasize that
there was no real conflict between a belief in selection and a belief
in an inheritance of acquired characters-for either Weismann or
Darwin. Both believed from the beginning in the overwhelming
importance of natural selection as the mechanism responsible for
the production of adaptation, but they desperately needed to find
some mechanism that would produce the variation needed for the
operation of natural selection. It was the task of the inheritance of
acquired characters to supply at least part of this variation. That
both authors made use of this factor was understandable, for a
belief in an inheritance of acquired characters was virtually uni­
versal until the beginning of the 1 88os. This was as true for folk­
lore as for science. The few dissenters (Galton, perhaps His)
spoke with muted voices and were not heard.
In view of Weismann's repeated Lamarckian statements in
1 88 1 , it is rather surprising how' sweeping his repudiation of La-

• 1 1 9 .
 A Hard Look a t Soft In heritance

marckis m was i n his 1 8 8 3 lecture " O n Heredity. " His refu tation
of Lamarckian claims is S O• oroad, and his Darwinian in terp reta­
tion of the n u merous cases that had previously been cited as
p roofs for an inheritance of acquired charact ers is so wel l thought
out, that one is almost forced to the conclusion that Weis mann
must h ave been thinking about this problem fo r many years . De­
s pite its title, Weism ann's 1 8 8 3 pa per did not develop a theory of
inheritance but was devoted almost exclusively to the refutation
o f an inheritance o f acquired chara cters . His st rategy was re mark­
ably similar to D a rwin's w hen he refuted crea tionis1n : Weism ann
took up one case a fter another that simply could not be expl ained
b y " use and disuse" and o ther Lamarckian mechanis n1 s . How can
the nu merous s pecial ada ptations of the worker and soldier cas tes
of ants be inherited by use, when these cas tes do not reproduce?
How can habits beco me ins tincts through use, when a particular
instinct is pra cti ced only once in the whole life of the individual ,
as is so o ften the case of rep roductive instincts among in sects?
How can the external structure of insects be n1 odifi ed b y use and
disuse, when the chitinous s keleton is laid down during the pupal
s tage and never changes afterward? To a modern person , fully
convinced of the impossibility of an inheritance o f acquired char­
acters , Weism ann's argu n1 ents seem n1ost persuasive . But in
Weismann's time the belief in the Lamarckian p rinciple was so
deepl y ingrained that only a minority were converted . Use and
disuse seemed a far more convincing explanation of the loss of
extren1 ities by snakes or of eyes by cave animals . I t was not un til
the evolutionary s ynthesis of the 1 940s that un reserved selection­
i s 1n was n1 o re or less univers ally ado pted by biologists ; but the
con clusive refu tation of the p rinciple of the inheritance of ac­
quired characters wa s no t achieved until the 1 9 5 0s , through the
so-called cen tra l do g m a of molecular biology, which stated that
no inforn1ation con tained in the properties of the son1 a tic p ro­
tein s could be trans ferred to the nu cleic acids of O N A .
Weis 1n ann's strategy was to sho\v not only that an inheritance
of acquired cha racters encounters fo rmidable difficulties but also
that cases cited in its favo r could be explained quite \vell through

· 1 20 •
 A Ha rd Look at Soft Inheritance

the theory of natural selection. A structure that is used a great

deal in an individual's lifetime is of course also exposed to strong
. selection forces. If the organ is inferior in a particular individual,
its owner will be handicapped in the struggle for existence.
Therefore, "the improvement of an organ in the course of gener­
ations is not the result of a summation of the result of practice of
individual lives, but of the summation of favorable genetic fac­
tors" ( 1 883 : 26).
Weismann stated repeatedly that he was inclined to apply the
principle of selection far more consistently than Darwin himself.
The fact that in ducks and other domestic fowl the wings have
somewhat degenerated while the legs have become stronger than
in their wild ancestors was explained by Darwin as the result of
use and disuse. Weismann quite rightly pointed out �hat this can
be explained even better by assun1ing that natural selection was
the cause of this change in proportions.
It is rather ironical that natural selection is attacked in the cur­
rent evolutionary literature not so much for its inability to ex­
plain certain adaptations or other evolutionary developments as
for being a principle so successful that anything could be ex­
plained by natural selection-and therefore, to use Karl Popper's
language, that it would be impossible ever to refute any evolu­
tionary explanation based on the principle of natural selection.
This was not the situation in Weismann's day, when people had
not yet become accustomed to thinking in terms of natural selec­
tion and when they were far more comfortable explaining evolu­
tionary developments in terms of an inheritance of acquired char­
acters.
Weismann was in his mid-forties when he shifted from La­
marckism to an uncompromising selectionism. An important
role in his conversion was undoubtedly played by his own obser­
vation-and that of various cytologists and embryologists-that
the future germ cells in various types of invertebrates are set aside
after the first mitotic divisions of the developing embryo and no
longer have any physiological connection with the body cells
(Churchill 1 9 85) .

• 121 •
 A Hard Look a t Soft Inheritance

This observation led Weismann .. "

in 1 8 8 5 to his theory of the
" continuity of the germ pbsm," which states that the " germ
track" is separate from the body (soma) track from the very be­
ginning , and thus nothing that happens to the soma ca n be com­
municated to the germ cells and their nuclei . In this early theory
the separation was between germ cells and body cells . Even
though such a separation may occur in certain organisms during
the earlies t cell divisions of development, in most organisms
(particularly in plants) many if not most somatic tissues are able
to produce germ cells . For this reason Weismann replaced germ
cells by germ plasm in his later publications . The strict separation
we now make of the DNA program of the nucleu s from the pro­
teins in the cytoplasm of each cell reflects Weismann's early in­
sight (Churchill 1 9 8 5 ) .
Weismann was correct in concluding that the germinal material
is something entirely different from the body substance and " that
the differentiation of the body cells is not acquired by them di­
rectly, but that it was prepared by changes in the molecular struc­
tu re of the germ cell " ( 1 8 8 3: 1 4) .
Weismann came close to the concept of a genetic program
when he said that we will come to the right conclusions about
development " if we consider all processes of differentiation that
occur in the course of ontogeny as controlled by the chemical and
physical molecular structure of the germ cell " ( 1 8 8 3: 1 8) .

The S ignificance o f Sex and Genetic Reco mbination

The refutation of an inheritance of acquired characters seemingly
left a serious void in evolutionary theory. Weismann fully under­
stood that he had to find a new mechanism for the production of
genetic variability. His solution was this: " I believe that such a
source is to be looked for in . . . sexual reproduction . . . Two
groups of hereditary tendencies are, as it were , combined. I re­
gard this combination as the cause of hereditary individual char­
acters , and I believe that the production of such characters is the
true significance of [sexual] reproduction" ( 1 8 8 6 : 272) . Indeed,

· 1 22 •
 A Hard Look at Soft Inheritance

this p rocess of genetic reco mbination through sexual reproduc­

tion was recognized by Weismann as being one of the most im­
portant pro cesses in evolution , and to it he devoted an en tire long
essay ( 1 8 86) .
His conclusion was in total opposition to prevailing ideas .
Since blending inheri tance was widely accepted in 1 8 80, even by
D arwin (Mayr 1 9 8 2 : 779 -78 1 ) , who si multaneously believed in
particulate inheritance, sexual reproduction was credited with as­
suring the uniformity of species . That Weis mann could put forth
such revolutionary claims was made possible by the discovery of
van Beneden ( 1 8 8 3 ) and other cytologists that m aternal and pa­
ternal chro moso mes do not fuse during fertilization but merely
reestablish the diploidy o f the zygote. What sexual reproduction
thus a chieves is not a homogenization of the parental characters
but their reco mbination . Genetic reco mbination together with
natura l s election can thus bring together previously separate and
independent characteris tics that greatly improve the selection
value of their bearers . Concerning the origin of sexuality, Weis­
m ann's s tatements a re vague, i f not teleological .
To a rem a rkable extent Weis mann was aware also of the draw­
backs of sexuality. He des cribes the distinct advantage by which
the tem porary abandon ment of sexual rep roduction allows cer­
tain anim als such as aphids and cladocerans " a much more rapid
increase in the nu mber of individuals . . . in a given time"
( 1 8 8 6 : 2 8 9) . However, this is only a tem porary advantage-a con­
clusion confirmed, according to Weismann , by the fact that
" whole groups of purely parthenogenetic species or genera are
never met with " ( 1 8 8 6 : 290) . Although this statement is no longer
literall y t rue, we know of only a single higher taxon of animals ,
the bdelloid rotifers , in which all species are asexual . All other
asexually rep roducing groups of animals s eem to become extinct
sooner or later.
The overw helming importance of genetic reco mbination was
almost totally ignored by most Mendelians , owing to their
stron gly reductionist position . The broad recognition of reco m­
bination in the evolutionary process had to await the evolutionary

• 1 23 •
 A Ha rd Look at Soft Inheritance

synthesis . Those m athematical.'

population geneticists who em-
phasized the gene as the target of selection were particularly slow
to appreciate fully the role of reco mbination . I t is today q u ite
evident that what is o f particular i m portance in evolution is not
so m uch allelic interactions as interactions among different loci
and different chromoso mes , which are cons tantly changed by ge­
netic recom binatio n . Weis m ann's ( 1 89 1 ) cha mpionship of a mp hi­
mixis, as he called it, is one of his mos t important cont ribu tions
to evolutionary biology. Weism ann's explanation of the evolu­
tionary significance of sexu al reprodu ction has been challenged
in recent years b y a num ber of au thors , but no con sensus has yet
been reached as to a poss ible alternative explanation .

The Sou rce of New Genetic Varia tion

The mixing of the genetic fa ctors of both parents p rodu ces an
almost unlimited supply of genetica ll y new in dividuals in every
genera tion , but this process consists only in the in termingling of
already exis tin g variations . The ori gin of en tirely new genetic
fa ctor� re n1 a ins unexplained . The origin of true genetic novel ty
beca me a crucial p roblem fo r anyone rejecting an inheritance of
acquired characters , that is, a tran sfer of new somatic characters
to the gern1 pla s n1 . Weismann fully realized the seriousness of this
p roblem and struggled w ith it from 1 8 8 2 until fa r into the 1 8 90s .
He repeatedly s tated tha t organis ms normally give rise "only to
exact cop ies of themselves " ( 1 8 8 2 : 679, 6 8 2) . Furthermo re , natu­
ral selection is bound eventually to exhaust the supply of genetic
variants available through reco mbination . Thu s , Weis mann was
forced to come u p with a new solution ; and fo r thi s , he was ill
equipped (May r, 1 9 8 8 ) .
A n1ech anistic conception of causation co n1 pletely don1inated
Wei s rnann's thinking about the source of genetic va riation . When ,
prior to 1 8 8 3 , Wei sm ann gave up his belief in an inheritance of
acquired characters , he had to find an en tirel y new expla nation
for the origin of genetic variation . Being involved at that ti n1e in
a stru g gle with Naegeli , Kolliker, Eimer, and others about the

. 1 24 •
 A Hard Look at Soft Inheritance

exis tence o f in ternal forces of evolution , Weismann was unable to

adopt a process corres ponding to what we now call " s pon taneous
mutation . " Would not such a process have to be caused by inter­
nal forces ? This would o pen the doo r to what he called " meta­
physical p rinciples . " He wanted to explain everything " mechani­
cally, " and his con cept of mechanical was rather classical ,
i m plying the direct action of visible forces such as climate, nutri­
tion , and the like. I nstead , he postulated a nu1nber o f other pro­
cesses in the germ line, none of which tu rned ou t to be correct .
Ultimately all these explanations were based on the idea of mul­
tiple replicas of genetic determinants in the germ plas n1 and on
quantitati ve shifts in the relative number of these various ele­
ments .
The vigorous clai ms of Herbert S pencer and other neo­
Lamarckians that random varia tion was insufficient to supply the
needed material for the exercise of na tural selection left their
m a rk on Weismann's thinkin g . He eventually agreed with the La­
marckians that a sim ple Darwinian selection of individuals is not
sufficient to explain all the pheno mena of evolution ( 1 896: 5 9) , for
instance , the continuing reduction of vestigial organs ( 1 896 : 24) .
Finally, Weis mann admitted tha t chance alone could not produce
the right variation in the right species at the right time. When
perfect adaptation is achieved , " chance is ou t of the ques tion . The
variations which are su pplied to the natural selection of individ­
uals must have been produced [ within the germ] by the principle
of the su rvival o f the fi ttes t " ( I 896 : 46) . And this led Weismann to
propose his hypothesis o f germinal selection .
This is not the place to analyze the rationale of Weismann's
theory o f inheri tance, but one of its as pects must be mentioned
because it vitally a ffected Weism ann's evolutionary thinkin g . As
part o f the dominating influence of physicalism in nineteen th­
century s cience, quality was considered an unscientific concept .
Seemingly qualitative differences occurring in evolution had t o be
converted to quantitative differen ces . And this is precisely what
Weis m ann believed his theory o f inheritance could do . I t permit­
ted him to show that " all variation is in the last analysis quantita-

• 1 25 •
 A Hard Look a t Soft Inheritance

tive , consisting of an increas�, or decrease of the living particles or

of their constituents , the molecules " ( 1 904 2 : 1 2 8) . These changes
do not occur spontaneously but are always caused by external
factors , either by the differential nutrition of various determi­
nants within the germ plasm or by the environment.
Weismann apparently never realized that, notwithstanding the
terminology, his was no longer a selection theory. Indeed, it ap­
proached Geoffroyism, direct germinal induction by the environ­
ment, rather dangerously. Even thou gh Weismann continued to
deny emphatically any ability of the soma (phenotype) to affect
the germ plasm, in his theory of induced germinal selection he
admitted a direct effect of the environment on the gern1 plasm.
The most fundamental component of any theory of inheritance
is insistence on the basic constancy of the genetic material . Yet
the theory of germinal selection abandoned a constancy of the
underlying genetic elements . It seems that Weismann was forced
into this change of mind by the claims of various Lama rckians
that exposing the pupae of certain Lepidoptera to heat or cold
shocks had not only changed the coloration of the emerging but­
terfly but also that of some of its untreated descendants ( 1 904
2 : 2 3 0-23 1 ) . These results are now known to have rested on faulty
experimentation, and it is an irony of fate that they caused Weis­
mann to accept induced germinal selection and thus needlessly
undermined the consistency of his theory of inheritance . It is
worth noting that Darwin proposed his ill-fated theory of pan­
genesis also quite needlessly, because it was expressly proposed
to explain the effects of use and disuse, which one year after Dar­
win's death were shown to be illusory (Weismann 1 8 8 3 ) .
Weismann's theory of germinal selection was full of internal
contradictions and was aln1ost unanimously rej ected by his con­
te1nporaries , although he pleaded for it once more as late as 1 909
(pp. 3 6 -3 7) , as we 11 as in the third edition of his Vo rtrt({? e ( 1 9 1 o) .
All possible support for it had by that time been swept a\vay by
the acceptance of Mendelian inheritance . This included major
conceptual shifts , among them the acceptance of spontaneous
mutations and of qualitative genetic changes .

• 1 26 •
 A Hard Look at Soft l11lz erit a! 1ce

Weismann retained an unsha ken faith in Darwinian natural se­

lection all his life, in spite of the vicious attacks of l)riesch and
other experimental zoologists. When the fiftieth anniversary of
the publication of the Orziin of Species was celebrated in 1 909 ,
Weismann published a strong n1anifesto on the power of natural
selection. Such a declaration required considerable faith and cour­
age, since natural selection at that time was at the lowest point of
its scientific acceptance, owing to the attacks of de Vries, Bate­
son, Johannsen, and other Mendelians.

The Herita g e of Weis mann's Ideas

There can be little doubt that, after Darwin, no biologist in the
nineteenth century had as great an impact on evolutionary think­
ing as Weismann. A combination of characteristics enabled Weis­
mann to exercise this role. He had, on the one hand, a powerful
analytic a bility; he could build a logical argument step by step.
On the other hand, like D arwin, he had an extraordinary facility
for constructing hypotheses. This was by no means appreciated
in the nineteenth century; and, like Darwin, Weismann was ridi­
culed for his speculations. In view of the paucity of available facts
on inheritance, some of his speculations would be considered
rather daring even by a modern biologist who is constantly en­
couraged to try his hand at model building. When one reads the
attacks by Oscar Hertwig, Wolff, Driesch, and others, one real­
izes how outraged his contemporaries were by Weismann's theo­
nz1ng.
What Weismann did, he did for good reason. He was aware of
the intellectual poverty of the inductionism then dominant in
Germany. " The time in which men believed that science could be
advanced by the mere collection of facts has long passed away"
( 1 8 86 : 29 5 ) . Surely, it had passed away for Weismann, but most of
his contemporaries had not yet perceived the message. Until Dar­
win, Haeckel , Weismann, and a few other theorizers had begun
to develop a conceptual framework of biology, "the investigation
of mere details had led to a state of intellectual shortsightedness,

• 1 27 •
 A Hard Look at S o_ft In heritance

interest being shown only for that which was immediately in

view. Immense numbers oftdetailed facts were thus accumulated,
but . . . the intellectual bond which should have bound then1 to­
gether was wanting" ( 1 8 82:xv). Weis1nann again and again em­
phasized that his hypotheses were not the ultimate truth but were
proposed as heuristic devices. Of one of his theories he said,
"Even if it should be later necessary to abandon this theory,
nevertheless it seems to me to be a necessary stepping-stone in
the development of our understanding, it was absolutely neces­
sary that it had to be proposed, and it must be carefully analyzed,
regardless of whether in the future it will be found to be correct
or false" ( 1 8 85 : 1 7). This is a clear statement of his hypothetico­
deductive scientific philosophy.
To sum up, then, Weisn1ann's 1najor contributions to biological
thought were:
( 1 ) Defense o_( n a tu ra l selection . Fron1 about 1 890 to 1 9 1 o, Dar­
win's theory was threatened to such an extent by various oppos­
ing theories that it was in danger of going under. Wcisn1ann's un­
deviating support of natural selection at this tin1e was a major
contribution to the emergence of a strengthened Darwinism.
Weisn1ann helped to develop a methodology for the analysis of
natural selection, showing that it would pern1it the making of
predictions that would be confirmed if natural selection did in­
deed operate. He showed that the reduction or loss of structures,
which greatly disturbed some of his contemporaries, could be
explained as being caused by a relaxation of selection pressure.
Weismann was one of the first to test selection and environ­
mental influences by experin1ent. For instance, he exposed cater­
pillars of different colorations to potential predators on differ­
ently colored substrates. In other experiments he tested the effect
on the color of butterflies of the ten1perature at which the pupa
\Vas kept.
Perhaps even more effective than his evidence in favor of selec­
tionisn1 were his arguments against orthogenesis, saltationisn1,
and La1narckism, the three theories con1peting \Vith selection
(Bowler 1 983). He was particularly convincing in his argu111ent
in favor of the gradual nature of evolutionary change.

• 1 28 •
 A Hard Look at Soft ln lzl'rita11ce

(2 ) Re_futation o_f the theory o_f the inh eritance of acquired characters .
Through his s weepin g rej ection of an in heri tance of acquired
cha ra cters Wei s n1 ann es tablished a new version of Darwini s n1 .
The inheritan ce of acquired ch aracters never regained fu ll credi­
bility a fter Wei s rnann's a t tack in 1 8 8 3 . Wei s n1ann s u pported his
case b y three lines o f evidence : there i s n o cytological n1 echanism
that could effect s u ch a t ransfer fro n1 son1 a to gern1 pla s n1 ; there
a re m an y a d aptations that cou l d not have been acquired by such
an inheritance ( fo r exan1 ple, the soldier caste of ants and ter­
m i tes) ; and a l l repu ted cases of inheri tan ce of acqu ired cha racters
can be explained b y selection . Even though Weism ann was occa­
s io na l l y wrong in detail , he was right in prin ciple , and the basic
Wei s m an nian thou g h t is now articula ted in the so-called cen tral
dog ma of m olecular biology.
( 3 ) Finn establish men t o_f particu late inheritance. I f there were
blending inh eritance (at the gene level) , very differen t laws of ge­
netics (Galto n) wou l d be valid than if inheritance were p articu­
late . Fu rtherm o re, if each a ct of fertilization (zygote fo rmation)
con sists in the co mbining (ra ther than fu sion) of the paternal and
m a terna l gen o m es , then this must be com pensa ted by a redu ction
division, Wei s m ann h ypothesized (Churchill 1 96 8 ; 1 979; Farley
1 9 8 2) . These postu lates laid the fo undation fo r Mendelian genet­
ics (as clearly stated by C o rrens) , and Mendelian genetics in its
turn validated Wei s m ann's t heo ries .
(4) Reco�nition of the i1nportance o_f sexua l reproduction as a source o_f
genetic variation . The i m portan ce of genetic reco mbination de­
pends on particulate in heritance . I t wa s Wei s m ann who first real­
ized the vast i m po rtance of sexual repro du ction as a mechanism
fo r the p ro duction o f a l most unli mited genetic reco 1nbination
(Churchill 1 979) . A l though this fa cto r wa s rather neglected in the
m u tationist heyday of M en delism , it wa s revived fro m the 1 9 3 0s
on . Weis m an n , like the m o dern evolutionis ts , wa s a true fo llower
of D a rwin , for whom also the individual was the target of selec­
tion . Natu ra l selection wo uld he hel p les s if there were not an in­
exh austi b le s u p p l y of genetic varia tion in the form of uniq uely
differen t individual s .
( 5 ) Constraints on na tural selectio n . A s discussed above , Weis-

• 1 29 .
 A Hard Look a t Soft In heritan ce

mann strongly emphasized that there are severe developmental

and other constraints on tht"'.power of selection.
(6) Mosaic evolution . Weismann emphasized again and again that
not only different components of the phenotype but also different
stages in the life cycle vary in their rate of evolution. For instance,
in b utterflies the caterpillars often evolve faster and along entirely
different lines from the imagos ( 1 8 8 2 : 4 3 2) . As a result, a classifi­
cation based on larval characters is not at all the same as one based
on imagos . Weismann lists ins tances of mosaic evolu tion for
many groups of insects ( 1 8 82 : 4 8 1-50 1 ) . In mos t cases it results
from special adaptations of the larval stages . Indeed, Weismann
offers a long list of evolutionary " incongruences " discovered
when he compared either larval and adult stages or the corre­
sponding stages of more closely or more distantly related higher
taxa ( 1 8 8 2 : 502-5 1 9) .
(7) Cohesion of the genot ype. Weismann stated repeatedly that ,
mosaic evolu tion notwithstanding , there are limits to the inde­
pendence of different components of the genotype. Organisms
must evolve more or less harmoniously, and a selection pressure
on one organ very often results in a selection pres su re on some
other structure. Or a change in behavior results in the necessity
for modification of a structu re. In other words, it is the genotype
as a whole that responds to the forces of natural selection, a con­
sideration often ignored during the reductionist period of math­
ematical genetics .
More broadly, in an age of inductionism it was very important
that there was someone who had the courage to speculate. Weis­
mann pointed ou t significant problems and unanswered ques­
tions , even in cases where he himself was unable to find the right
solution.
Probably no one in the latter part of the nineteenth century
comprehended the basic thesis of Darwinism better than Weis­
n1ann. He was the only one who understood the overwhelming
role of natural selection. And he realized that the source of genetic
variation was the great unknown in the process of selection and
that a detailed theory of inheritance was the need of the hour.

• 130 •
 A Hard Look at Soft Inherita_nce

Even though he hi mself failed to 1neet that need , the in tellectual

preparation he gave to the a rea enabled the Mendelian theory to
prosper. Even more than in his own lifetime, Weismann is today
considered one o f the very few truly outs tanding evolu tionary
biologis ts .

. 131 •
 CHAPTER NINE
.. '
-----------·-·--• -------------

Geneticists and N aturalists

Reach a Consensus :
The Second D arwini an Revolution

/\ o vANCE I N SCI ENCE is rarely s teady and regular. Nor can

..fi. it necessarily be des cribed in Thomas Ku hn's tern1 s as a
series of revolutions separa ted by long periods of steadily pro­
gressing normal s cience . Rather, when we study particular scien­
tific disciplines we observe great irregulari ties : theo ries bcco n1c
fas hionable, others fa ll in to eclipse; son1e fields enj oy con sider­
able consensus a 1n ong thei r active wo rkers , other fields arc s plit
into several ca rn ps of specialists furiou sly feudin g with one an­
other. This l a tter description applies wel l to evolutionary biology
between 1 8 5 9 and about 1 940 .
The oppos ition to natural selection con tinued unabated fo r
some eighty years after the publication of the Origi11 . Except fo r
a few naturalists , there was hardly a single biologist, and certainly
not a s ingle ex peri mental biologist, who ado pted natural selec­
tion as the exclusive ca use of adaptation . One wo uld have
thou ght that the rediscovery of Mendel 's laws in 1 900 would have
brou ght about an i m mediate change in the attitude toward na tu­
ral selection , but this was not the case. The findings of genetics
n1 ade it now q uite clear that the genetic 1n aterial wa s particulate
(hence inheritance could not be blen din g) , and likewise that in­
heritance was hard (that i s , not permi tting any in heri tance of ac­
quired characters) . Yet the leadin g Mendelians-Bateson , de
Vries , and Johannsen- did not adopt natural selection . They as­
cri bed evolu tion ary change ins tead to n1 utation pressure .
B y the 1 9 20s n1 ost studen ts o f evolu tion belonged t o one o f
th ree biolo gical disciplines : genetics , systern a tics , or paleonto l-

. 132 •
 Geneticists and Na tu ra l ists Reach a Consensus

ogy. They differed in their interests and in the kind of knowledge

they had. The naturalists (both the systematists and the paleon­
tologists) were not sufficiently familiar with the advances in ge­
netics made after 1 9 1 o, and thus they were arguing against the
erroneous evolutionary concepts of the Mendelians as though
this were still the viewpoint of genetics. The geneticists, in turn,
ignored the rich literature on geographic variation and speciation;
consequently, nothing in the evolutionary writings of T. H. Mor­
gan, H. J. Muller, R. A. Fisher, or J. B. S. Haldane could explain
the multiplication of species, the origin of higher taxa, or the ori­
gin of evolutionary novelties. Furthern1ore, the two groups dealt
with different hierarchical levels: the geneticists with intrapopu­
lational variation at the gene level, the naturalists with the geo­
graphic variation of populations and with species. When geneti­
cists and paleontologists, or geneticists and taxonomists, had
joint meetings in that period, their respective backgrounds were
so different that they were seemingly unable to communicate
with one another.
Yet during the 1 920s the foundation was laid for an eventual
consensus. Not only were blending as well as soft inheritance def­
initely refuted by the geneticists during this period, but the oc­
currence of spontaneous mutations was firmly established, some­
thing the deterministic physicalists of the nineteenth century had
been unable to accept. Also, Morgan and his school as well as
Edward East and Erwin Baur found that most mutations had
only very small effects on the phenotype and were not at all like
the large mutations envisaged by the early Mendelians. In this
period the difference between genotype and phenotype was clar­
ified, and it was understood that what is selected are whole gen­
otypes, not individual genes. Therefore, genetic recombination
rather than mutation was seen as the immediate source of the ge­
netic variation available for selection. The way selection acts in a
population was understood much better when it was connected
with these improvements of understanding.
Rather unexpectedly, within a few years, a wide-reaching con­
sensus among genet1c1sts, systemattsts, and paleontologists was

• 1 33 .
 Gen eticists and Natu ra lists Reach a Consensus

rea ched . The term "evolu tionary s y n thesis " was in trodu ced b y
Ju lian H u xley ( 1 94 2 ) t o designate t h e acceptance of a unified evo-
1 u tion ary theory b y the previousl y feuding camps of evolu tion­
ists . H ow could this unexpected es tablish men t of seen1ing ag ree­
men t h ave come to pass so su ddenl y ? Two wo rkshops were
organized in 1 974 in which an answer to this ques tion \Vas sought
(Mayr and Provine 1 9 80) . I t beca me clea r that the synt hes is o f the
opposing viewpoints was n1 ade possible \vhen a n u 1n ber of ta x­
ono1n is ts- Sergei C h etverikov , Theodosius Dobzhansky, E. B .
Ford , Bernhard Ren sc h , and I -becan1e acquain ted \V ith post­
M endelian genetics (that is , popula tion genetics) and devel oped
an u p-to-date l) arwinisn1 that co n1bined the bes t ele111 cn ts o f
both gen etics a n d sys ten1atics . Fu rther more , George Gaylord
Sim pson was res ponsi ble fo r bringin g paleontology and 1n acro­
evolution into the s y n thesis . He wa s able to sho\v that the phe­
no men a stu died b y the paleon tologist-that is , n1 acroevolu tion ,
o r evolu tion above the species level -is in every res pect con sist­
ent with the findings o f n1 odern genetics and w i th the basic con­
cepts of D a rwinism . The s a 1ne was shown in dependen tl y in Ger­
many b y Rens ch and i n A m erica , fo r plan ts , b y G . L . Stebbi n s .
The ne\v con sensus wa s heral ded and pro n1 o tcd b y Dobzhan­
sk y 's Genetics and tlzc Origin o_f Species ( 1 9 3 7) , followed by J . H u x ­
ley 's Evolution: Tlz e Modern Syn th esis ( 1 942) , May r's Systematics
and th e Or(gin of Species ( 1 942) , Sin1 p son's Tempo and Mode in Evo­
lution ( 1 944) , Rensch 's Neuere Prohlem e dcr A bsta,nnz ungslch re
( 1 94 7) , and S tebbins 's Varia tion and Evolu tion in Plants ( 1 9 5 0) .
These autho rs have often been referred to as the architects o f the
evolutionary s y n thesis . This is a somewhat arbitrary selection ;
one could have incl uded among the architects also some evol u­
tio nists w h o pu blis hed prior to 1 9 3 7 , su ch as Chetverikov , S u m­
ner, St resen1a nn , Fisher, Haldane, and Wrigh t .

The Nature of the Evolutionary S ynthes is

What occu rred during the period fro n1 1 9 3 6 to 1 9 5 0 , \v hen the
s y n thesis took place , was not a s c ien tific revolu tion ; rather it \Vas

• 134 .
 Geneticists and Na tu ralists Reach a Consensus

a unification of a previously badly split fiel d . The evolu tionary

synthesis is important because it has taught us how such a unifi­
cation may take place : not so much by any revolutionary new
concepts as by a p rocess of house cleaning , by the final rej ection
of various erroneous theories and beliefs that had been res pon­
sible for the previous dissension . Am ong the constructive
achievements o f the synthesis was the finding of a com n1on lan­
guage among the pa rticipating fields and a cl arifica tion of n1any
aspects of evolution and its underlying concep ts .
The period of the synthesis was not one of grea t innovations
but rather of mutual education . Naturalists who had not known
it befo re learned from the geneticists that inheritance is al ways
hard, never soft . There can be no heritable influence of the envi­
ron ment , no inheritance o f acquired characters . Weis mann's the­
sis was finally adopted universally more than fifty years a fter it
had fi rst been p roposed . Another finding of genetics , its particu­
late chara cter, was also finally universally adopted . Many natural­
ists up to that time had divided characters into Mendeli an (partic­
ulate) ones , which they considered evolutionarily unimportan t ,
and gradual o r blendin g ones , which , following Darwin , they
considered to be the true material of evolution .
Acceptance o f these two findings of genetics helped in the ref­
utation o f the three m aj or evolutionary theories that had com­
peted with natu ral selection since the publication of the Origin
(Bowler 1 98 3 ) . These theories , as we have seen , were ( 1 ) neo­
Lamarckism (the inheritance of acquired ch aracters) and other
forms of soft inheritance; (2) autogenetic theories based on the
belief in a built-in drive toward evolutionary progress ( ortho­
genesis, nomogenesis, aristogenesis , omega principle) ; ( 3 ) and
saltational theories of evolution , which pos tulated the sudden
emergen ce of drastically new life forms (de Vriesian mutations) .
Perhaps no author contributed more to the refutation of these
three theories than G . G . S im pson, whose Tempo and Mode ( 1 944)
and Mean ing of Evolu tion ( 1 949) consist in large part of evidence
dis p roving them .
But the s ynthesis was not merely the general acceptance by the

• 1 35 .
 Geneticists and Natu ralists Reach a Consensus

naturalists of the principles of theoretical population genetics. To

understand the achievements of the synthesis, one must appre­
ciate how typological and saltationist the evolutionary views of
the original Mendelians were (Provine I 97 1 ) . It is a Whiggish
misrepresentation of history to equate Mendelism with genetics .
To be sure, the two agreed in their rej ection of soft and blending
inheritance . On the other hand, the evolutionary views of Bate­
son, de Vries, and Johannsen were rejected during the synthesis
(Mayr and Provine 1 9 80) . Indeed, neo-Lamarckism seemed to
explain evolution better than did the saltationist theories of the
Mendelians.
A major achievement of the synthesis, then, was to develop a
unified view on the nature of genetic change. Darwin , accepting
universal opinion on this subj ect, had thought there were two
kinds of variation: drastic ones, often referred to as sports, and
small ones, represented by gradual or quantitative variation. For
Darwin, it was gradual variation that was important in evolution.
By contrast, the Mendelians insisted that new species originated
through drastic mutations. Genetic research by Nilsson-Ehle,
East, Castle, and Morgan during the first third of the twentieth
century showed clearly that drastic and minutely differing var­
iants were only extremes of a continuously varying spectru n1 ,
and that the same genetic mechanism was involved in n1utations
of all degrees of difference.
This finding had a number of important consequences. It per­
mitted a reconciliation between the Mendelians and those who
studied quantitative inheritance . It also permitted the building of
a bridge between n1icro- and macroevolution. Most irnportant,
it refuted the credo of essentialism. There is no uniforn1 species
essence, but rather each individual has a highly heterogeneous
genotype (which varies fron1 individual to individual) . The belief
in two kinds of variation was still widespread until the 1 93 0s, but
the new way of interpreting genetic variation became con1pletely
victorious during the synthesis (Sapp 1 98 7) .
The realization that gradual variation could be explained in
tenns of Mendelian (particulate) inheritance also led to the end of

. 136 .
 Gen eticists and NatH ralists R each a ConsensHs
.,

any belief in so-called blending inheritance . This was helped by

the clear recognition of a difference between genotype and phe­
notype, as was shown by Nilsson-Ehle, East, and others; a con1-
plete " blending'' of characters of the phenotype was possible in
spite of the discrete particulateness of the underlying genetic fac­
tors.

C o n tributions by the Naturalists

Because of these advances in genetics, it is therefore son1eti111es
asserted that the evolutionary synthesis was merely the applica­
tion of Mendelian inheritance to evolutionary biology. This for­
mulation ignores two important factors . One is that the geneti­
cists, particularly the experirnental and the mathematical ones,
had to acquire j ust as n1uch fro111 the naturalists as the naturalists
had to acquire from the geneticists. And second, the conceptual
framework of evolutionary biology was greatly enriched by con­
cepts and facts from natural history that were conspicuously ab­
sent in the writings of the geneticists.
For example, evolution, defined by the geneticists as "a change
of gene frequencies in populations," was visualized essentially as
a strictly temporal phenomenon. This is reflected in Muller's
statement: " Speciation represents no absolute stage in evolution,
but is gradually arrived at, and intergrades imperceptively into
racial differentiation beneath it and generic differentiation above "
(Muller 1 940: 2 5 8) . Throughout their writings the geneticists
concentrated almost completely on the temporal component in
evolution. The paleontologists, by necessity thinking in terms of
vertical sequences, likewise confined themselves to the study of
vertical evolution until they merged their thinking with the hori­
zontal tradition of the naturalists (Eldredge and Gould 1 972) .
One of the most important contributions of the naturalists, the
heirs of Darwin, was to bring geographical thinking into the syn­
thesis. The problem of the multiplication of species, the existence
of polytypic species, the biological species concept, the role of
species and speciation in macroevolution, and many other evolu-

• 137 .
 Gen eticists a nd Naturalists Reach a Consensus

tionary problems can be dealt with only by invoking geographi-

cal evolution. ·:'
The incorporation of the geographical dimension was of par­
ticular importance for the explanation of macroevolution. Pa­
leontologists had long been aware of a seeming contradiction be­
tween Darwin's postulate of gradualism, confirmed by the work
of population genetics , and the actual findings of paleontology.
Following phyletic lines through time seemed to reveal only min­
imal gradual changes but no clear evidence for any change of a
species into a different genus or for the gradual origin of an evo­
lutionary novelty. Anything truly novel always seemed to appear
quite ab ruptly in the fossil record. During the synthesis it became
clear that since new evolutionary departures seem to take place
almost invariably in localized isolated populations ,· it is not sur­
prising that the fossil record does not reflect these sequences . A
purely vertical approach is unable to resolve the seeming contra­
diction.
An equally important contribution made by the naturalists was
the introduction of population thinking into genetics . Mendelism
was strongly typological-the mutation versus the wild type.
And even later, in the thinking of many geneticists , not only
Morgan and Goldschmidt but even M uller in his search for the
perfect genotype, and R. A . Fisher, one can detect a strong es­
sentialistic component. Population thinking , with its emphasis
on the uniqueness of every individual in the population, was
brought into genetics by Chetverikov and his students (including
Timofeeff-Ressovsky) , by Dobzhans ky, and by Baur. With few
exceptions (for exan1ple, polyploidy) , every evolutionary phe­
nomenon is simultaneously a genetic phenomenon and a popula­
tional one.
Finally, the naturalists , or at least some of them, attempted to
replace the strictly reductionist formulation of most geneticists
by a more holistic approach. Evolution, they said, is not merely
a change in the frequency of genes in population� , as the reduc­
tionists asserted, but is at the same time a process relating to or-

. 1 38 .
 Geneticists and Natu ralists Reach a (;onsensus

gans , behaviors , and the in teractions of individu als and popula­

tions . In this holis tic attitude the naturalists agreed with the de­
velo pmental biologists .

The Triun1ph o f Natural Selection

The synthesis was a reaffirmation of the Darwinian formulation
that all adaptive evolutionary change is due to the directin g force
of natural selection on abundantly available variation . Today we
a re so com p letely used to the Darwinian formula tion that we are
apt to forget how different it is from the evolutionary explana­
tions of D a rwin's opponents .
The naturalists, in the tradition of Darwin , had been the
staunchest defenders of natural selection from the very begin­
ning , but like D arwin almost all of them tended to believe simul­
taneously in a certain a m ount of soft inheritance . Now that this
form of inheritance was decisively refuted, the stronges t support­
ers of natural selection were found among the naturalists .
There i s , as yet , no good history of the acceptance of natural
selection by the geneticists . C hetverikov , Timofeeff-Ressovsky,
and D obzhans k y got it fro m a strong Russian tradi tion (Adams
1 9 80) . In no o ther country did natural selection have as wide­
spread a bas e of s u pport as in Ru ssia. But for ideological and po­
litical reasons this highl y successful genetic establishment was
wiped out b y S talin's regime and in its place was ins talled the
charlatan Lysenko and his henchmen . In England there was a
strong selectionist tradition at O xford ( Lankester, Poulton , and
so on) , but the classical Mendelians had no use for selection , and
in the United States Morgan's thinking was very much in that
tradition . A second tradition in the United States , however­
cen tered in H arvard 's Bussey Institution (Castle, East, Wright) ­
ap parentl y adopted selection without reservation . In France ,
Lwoff at t h e lnstitu t Pasteur seems t o have been the first consist­
ent selectionist, followed by Ephrussi , L ' Heritier, and Teissier.
But even now strict selectionists seem to be in the minority

• 1 39 .
 Geneticists and Na tu ralists Reach a Consensus

among French biologists. In every country except the U SSR se-

.. '
lectionism gradually grew i.n strength in the years after the syn-
thesis.
It is understandable that in the early stages of the synthesis the
universal presence of natural selection should have been empha­
sized strongly, since a considerable number of Lamarckians still
existed among the older evolutionists. However, as soon as this
stage passed, there developed a trend toward recognition of other
factors. Where the modern biologist perhaps differs from Darwin
most is in assigning a far greater role to stochastic processes than
did Darwin or the early neo-Darwinians. Chance plays a role not
only during the first step of natural selection-the production of
new, genetically unique individuals through reco1nbination and
mutation-but also during the probabilistic process that deter­
mines the reproductive success of these individuals. All sorts of
constraints forever prevent the achievement of "perfection. " Even
though natural selection is indeed an optimization process, the
existence of numerous opposing influences n1akes optimality
quite unachievable.
The unification of evolutionary biology achieved by the syn­
thesis painted its picture in bold strokes: Gradual evolution is due
to the ordering of genetic variation by natural selection, and all
evolutionary phenomena can be explained in terms of the known
genetic n1echanisn1s. This was an extreme simplification, consid­
ering that processes in organismic biology are usually highly
complex, often involving several hierarchical levels and pluralistic
solutions. The task of evolutionary biology after the synthesis of
the 1 940s was to convert the coarse-grained theory of evolution
into a fine-grained, more realistic one. No longer constrained to
a defense of Darwinism, the followers of the evolutionary syn­
thesis, in the course of the more detailed analysis, began to tackle
differences that still existed, not only between the reductionist
tendencies of the geneticists and the organismic viewpoint of sys­
tematists and paleontologists but also concerning other aspects of
evolutionary theory.

• 1 4 () .
 C HAPT E R T E N
-------------•-------------

New Frontiers in
Evolutionary Biolog y

J
usT As IN THE DEcAoE after the rediscovery of Mendel 's
rules, since about 1 970 the claim has been n1ade increasingly
often that "Darwinisrn is dead." I shall not deal at all with the
attacks by creationists, based on ideological commitments, since
their argun1ents have been decisively refuted by Futuyma ( 1 983 ),
Kitcher ( 19 82), Montagu ( 1 9 83 ), Newell ( 1982), Ruse ( 1 9 82),
Young ( 1 9 85 ), and several other authors. Claims that Darwinism
is obsolete have been made in nun1erous articles and books also
by several nonbiologists, whose arguments, though nonreli­
gious, are based on such ignorance of evolutionary biology that
it is not worthwhile to provide references to their writings. More
disturbing are the similar, although somewhat more muted,
claims that have been made by some knowledgeable biologists­
even evolutionary biologists. These include Gould ( 1977; 19 80),
Eldredge ( 19 85 ), W hite ( 1 9 8 1 ), and Gutmann and Bonik ( 19 8 1 ).
Different critics have singled out different aspects of the synthesis
as particularly vulnerable. It has been claimed, for example, that:

( 1 ) The findings of molecular biology are incon1patible with Dar-

w1n1sm.
(2) The new research on speciation shows that other modes of spe­
ciation are more widespread and more i rnportant than geo­
graphic speciation, which the neo-Darwinians clain1 is the pre­
vailing mode.
( 3 ) Newly pro posed evolution ary theories, l ike punctu ationism, arc
in compatible with the synthetic theory.
(4) The synthetic theory, owing to its reductionist viewpoint, is un­
able to explain the role of development in evolution.
 New Fron tiers in Evolutionar y Biolog y

( 5 ) Even if one rejects the reductionist claim of the gene as the ta rget
of selection , Darwinism,. oy considering the individual the ta rget
of selection , is unable to explain phenomena at hierarchical levels
above the individual , that is , it is unable to explain macroevolu­
tion .
( 6) By adopting the " adap tationist program," and by neglecting sto­
chastic processes and cons traints on selection , particularly those
posed by development, the evolutionary synthesis paints a mis­
leading picture of evolutionary change .

These highly diverse criticis ms range from the extreme view

that D arwinism as a whole has been refuted to the milder op inion
that the s yn thesis is too na rrowly adaptationis t or that the con­
cept of s peciation has to be thorou ghly revised . One by one these
various criticis m s have been refuted b y A yala ( 1 9 8 1 ; 1 9 8 3 ) , S teb­
bins and A yala ( 1 9 8 1 ) , Grant ( 1 9 8 3 ) , Maynard S m ith ( 1 9 8 2 ;
1 98 3 ) , Mayr ( 1 9 8 4) , A . H uxley ( 1 9 8 2) , Levinton ( 1 9 8 8 ) , and
others .
The as pect o f almost a l l of these obj ections that I wish to ad­
d ress here is an evident misu nderstandin g o f the theory that
emerged fro m the evolutionary s ynthesi s . O ften this misunder­
s tanding a rises fro m the assumption that the most extreme reduc­
tionist version o f the synthesis , as represen ted b y s o me o f the
mathe m a tical population geneticists , is the basic dogma of the
syn thesis . When critics p ropose that the conclusions of the s yn­
thesis be repl aced by a more modern view of evolutio n , one finds
p ractically withou t exception that these supposedly novel views
have been m ain tained all along by Rensch, m ysel f, and several
other rep resentatives o f the s ynthes is . Almost a l l the critics o f the
synthesis , I a m sorry to s ay, have quite cons picuously m is rep re­
sen ted the views o f its leading spokes men .
To take one exan1 pl e , the critics con tin ually criticize the cl aim
that " D a rwinian evolution is due to the natura l selection o f ran­
d o n1 n1 utations . " This criticism co m pletely ignores the fa ct that
fro m D arwin o n to the 1 9 80s the indi vidual as a whole was con­
sidered the t a rget of selection fo r the organis n1ic biologis t s , and
therefo re reco mb ination and the stru ctu re of the genotype as a

• 1 42 •
 New Frontiers in Evolution ary ,B iology

whole were viewed as being far more im portant for evolution

than m utational even ts at individual loci . Fu rtherm ore, the critics
completel y misin terpret the word " random ." The term , when
a pplied to varia tion , n1eans that it is not in a res ponse to the needs
of the organis m .
O pponents o f the synthesis consisten tly con found three
schools of D arwinis m : ( 1 ) neo-D arwinism , a term coined by Ro­
manes in I 8 96 to designate " D arwinis m without an inheri tance
o f acquired characters " ; (2) early population genetics , a strongly
reductionist school that defined evolution as the modification of
gene frequencies by natura l selection ; and ( 3 ) the holistic branch
o f the s ynthesis, which continued the tradition s of Darwin and
the natu ralists while accepting the findings of genetics .
The thinking o f the reductionists was strongly influenced by
R. A . Fisher, and this school has therefore sometimes been des­
ignated as Fisherian D arwinis m . It is clearly the primary target of
the opponents of the s ynthesis, but these cri tics confuse matters
when t hey designate the reductionist s chool as neo-D arwinis m
or imply that it includes people like Huxley, Dobzhansky,
Wright, Rensch, or m yself, all of whom dis tinctly rej ected the
reductionist conclusions of the Fisherian school .
D a rwinis m is not a simple theory that is either true or false but
is rather a highly com plex research program that is being contin­
uously modified and improved . This was true before the synthe­
sis, and it continues to be true after the synthesis . Table 2 lists
many of the significant stages in the modification of Darwinism
that one might recognize. Yet recognizing such s eemingly dis­
con tinuous periods is in m any respects an artificial enterp rise . To
take but a few exa mples , the prevalence of particulate inheritance
was obvious after 1 8 86 but was not adopted until after 1 900; the
emphasis on the role of diversity in evolution was stressed by
naturalists from D a rwin on but was almost entirely ignored by
the Fisherians; the naturalis ts , for their part, rej ected the beanbag
genetics of the redu ctionists and during the post-synthesis period
con tinued their holis tic tradition of emphasizing the individual as
the target of selection . In short, each of these periods was heter-

• 1 43 .
 Ne w Frontiers in Evolu tionary Bio logy

ogeneous to some extent, owing to the diversity in the thinking

of different evolutionists . Most critics who have attempted to re­
fute the evolutionary synthesis have failed to recognize this diver­
sity of views and thus have succeeded in refuting only the reduc­
tionist fringe of the Darwinism camp. Their failure to appreciate
the complexity of the evolutionary synthesis has led them to
paint a picture of that period that is at best a caricatu re .
Another error made by most opponents of the synthesis is a
failure to differentiate between proximate and evolutionary cau­
sations . For Da rwin and all of his holistic followers , selection
starts at fertilization and continues through all emb ryonic and lar-

TABLE 2
Significant stages in the n1odification of Da rwinisn1 .

Date Stage Modifica tion

1 8 8 3 ; 1 8 86 Weismann's neo­ End of soft inheritance; diploidy

Darwinism and genetic reco 1nbina tion
recognized
1 900 Mendelism Genetic constancy accepted and
blending inheri tance rejected
1 9 1 8 - 1 93 3 Fisherism Evolution considered to be a matter
of gene frequencies and the force of
even s 1nall selection pressu res
1 93 6- 1 947 Evolutionary Population thinking en1phas ized ;
synthesis in terest in the evolu tion of di versity,
geographic speci ation, variable
evolu tionary rates
1 947- 1 970 Post-s ynthesis Individual in creasingly seen as
target of selectio n ; a more holistic
approach ; increa sed recogn ition of
chan ce and constraints
Punctuated I m portance of speciational evo lution
equilibria
Redis covery of I n1 po rtance of reproductive su cces s
se xual selection fo r select ion
 Ne w Fro n t iers in Ei 1 0/utionar y B,ioloJ! Y

va l s tages . A D a rwinian is truly puzzled when he rea ds in a cri­

tiq ue by an e111 b ryologist th a t " devel o p n1ent . . . co n1es to the
fore as a p roblem unintelligible within neo-Da rwinis n1 . " What
a spect o f develop men t is this author talk ing abou t? I f he is s peak­
ing o f the translation of the genetic p rogran1 in to molecular
chains o f even ts during on togeny, he i s talking about proxim ate
causa tion s . Their s tudy, indeed , has never been the j o b of the evo­
lutionary biolog i s t . But n1 any other as pects of develop ment do
raise qu estions con cerning evolutiona ry ca usa tions and these have
been o f in teres t to evolutionists from D a rwin on . They arc of
con cern to the evolu tionist, firs t , because each s tage of develop­
m ent is a ta rget o f selection , and particularl y so when the devel­
op mental s tages (larvae) a re free-livin g . Second , becau se em­
bryonic s t a ges m ay the n1sel ves serve as " soma tic programs " in
devel o pmen t (see bel ow) , such sta ges tend to beco me highly con­
served in evolu tion (for instance , the gill arch s tage of the tetra­
pod e mb ryo) . Such highly conserved s ta ges are often hel pful in
the reconstruc tion of the phylo geny (reca pitu lation) . No Dar­
winian will ever question the i m portance o f development in evo­
lutio n , but evolution ary in terpretation is cons tra ined by the ex­
tent to w hich the p roximate causations of develo pmen t have been
elucidated by the embryologists . To unde rtake a s tudy of such
proxi m a te causations is not the task of the evolutionis t .
C u riously, the obj ection i s sometimes raised tha t the evolu­
tionary s yn thesis can shed light neither on the level of the gene
nor on t rans-speci fi c levels because it i s concerned with indi vid­
uals (as t argets of selection) and with populations (as incipien t
species) . What is needed , i t is claimed, is a hierarchical approach
tha t i s not fo und in either neo-D arwinis m o r the evolu tionary
synthesis . Th a t this o bj ection is w ithou t basis has been shown by
Grant ( 1 9 8 3 : I 5 3 ) , S tebbins and A yala ( I 9 8 1 ) , and other defen ders
o f the synthesis . E ven S i m pson's interp reta tion o f evolution was
strong l y hierarchical (La porte 1 98 3 ) . No one has shown as clearl y
as M ayr ( 1 963 : 62 1 ) that the s pecies is the unit of action in ma cro­
evolution .
A n d finally, fo r White ( 1 9 8 1 ) , the s yn thesis wa s prema ture ,

• 1 45 •
 New Frontiers in Evolutionary Biology

"because there was no molecular biology at the time and both the
chemical nature of the hereditary material and the architecture of
eukaryote chromosomes were yet unknown." This is like saying
that the first Darwinian revolution was premature because genet­
ics had not yet been founded. Any scientific revolution or synthe­
sis has to accept all sorts of black boxes, for if one had to wait
until all black boxes are opened, one would never have any con­
ceptual advances.

The Evolutionary Synthesis as Unfinished Busines s

The healthy turmoil that currently characterizes evolutionary
biology should not be viewed as a death struggle but rather as the
sort of lively activity one will find in any healthy and advancing
branch of science. Yet it must be admitted that, even though the
refutation of the major anti-Darwinian theories during the syn­
thesis drastically narrowed down the variation of evolutionary
theory, some well-defined differences among the Darwinians still
existed into the post-synthesis period, and some of these differ­
ences are still with us fifty years later.
For the geneticists-or at least all those influenced by Muller,
Fisher, and Haldane-the gene continued to be the primary tar­
get of selection, and most genes were believed to have constant
fitness values. The whole problem of the origin of organic diver­
sity (that is, the multiplication of species) was minimized by this
school, if not ignored. Most of those who called themselves pop­
ulation geneticists worked with single, closed gene pools. Even
Wright did not come to grips with the problem of the multipli­
cation of species in his shifting-balance theory, nor with the ma­
croevolutionary problems generated by speciation.
By contrast, those who had come to evolutionary biology
from systematics or some other area of natural history considered
evolution to be a populational problem, and the whole (poten­
tially reproducing) individual to be the target of selection. For
them , the multiplication of species was the path\vay to the solu­
tion of problems of n1acroevolution. Despite their tendency to
 New Frontiers in Evolutionary I}iolo'-{? Y

think in terms of phenoty pes , they eventually came to view the

genotype as a systen1 of gene interaction-that is, they recog­
nized the cohesion of the genotype-and tended from the very
beginning to deal with evolution hiera rchically. Fo r Simpson ,
Rensch , Huxley, and M ayr, evolution was not a change in gene
frequencies but the twin processes of adaptive change and the ori­
gin o f diversity.
By no means are all current in tra-Darwinian cont roversies
remnants of the old geneticis t-versus-natu ralist feud. There are
also differences among the geneticis ts concerning the relative fre­
quency of neutral mutations and the amount of va riation due to
balan ce . A mong the paleontologists there are disputes as to
whether or not phyletic gradualism can give rise to higher taxa .
And among other evolutionists there are disagreements on the
validity o f g roup selection , the extent of adaptation, the role of
co mpetition , the frequency of sym pa tric speciation , how contin­
uous or punctuated evolution is, to what extent all co mponents
of the phenotype reflect ad hoc selection , what proportion of spe­
ciation is allopatric, what the target of selection is , how much
genetic variation is stored in populations , and to what extent the
new findings of molecular biology require a revision of current
theory. H owever, regardles s of ul timate outcome, none of these
disagreements affects any of the basic principles of Darwinism .
Although rarely com pletely wron g, the conclusions suppo rted
by the followers of the evolutionary synthesis were often inco m­
plete and rather sim plis tic. Two kinds of processes, in particular,
were o ften inadequately considered : ( 1 ) multiple simultaneous
causation s , and (2) pluralistic solution s . Let me give so me ex­
a m ples o f simultaneous causations . In all selection phenomena­
and selection is of course an antichance process-chance phe­
no mena also occur simultaneously. Or, to give another example,
speciation is never merely a matter of genes or chromosomes but
also of the nature and geography of the populations in which the
genetic changes occu r. Geography and the genetic changes of
populations affect the speciation process simultaneously.
Far more serious was a frequent neglect of the pluralism of evo-

• 147 •
 Ne w Fro n tiers in Evolu tio n a ry Biology

lutionary phenomen a . Darwin .. '

was well aware of the pluralis n1 of
evolutionary processes . Th� more one studies these processes ,
the more one is impressed by their diversity. Many con troversies
in evolutionary biology arise because of the inability of certain
authors to appreciate this diversity. There seem to be multiple
solu tions to almost any evolutionary challenge. During specia­
tion, pre-mating isolating mechanis ms originate first in son1e
groups of organisms; in others , post-mating mechanisms origi­
nate firs t . Sometimes geographic races are phenotypically as dis­
tinct as good species , yet not at all reprodu ctively isolated . On
the other hand, pheno typically indistinguishable spec ies (sibling
species) may be fully isolated reproductively. Some species are
extraordinarily young, having origin ated only 2 , 000 to 1 0 , 000
years ago , while others have not changed visibly in 1 0 to 5 0 mil­
lion yea rs . Polyploidy or asexual reproduction is i n1portan t in
some groups and totally absent in others . Chrom oson1al res truc­
turing seems to be an im portant co mponent of specia tion in some
groups, such as the morabine grasshoppers of Australia, but
seems not to occur in others . lnters pecific hybridiza tion is fre­
quent in some groups but is rare or absent in others . Some grou ps
speciate profusel y ; in others speciation seen1 s to be a ra re event.
Just because there is little gene flow in certain species , one can­
not conclude that gene flow is irrelevant in all species . In fact, the
amount of gene flow seen1s to differ greatly even in closely related
species . The absence of major genetic recon struction in many
founder populations does not prove that genetic revolutions can
never take place . Parapatry is frequent in groups in which post­
zygotic isola ting mechanisms evolve first when pre-zygotic ones
are still very incomplete; however, this does not j ustify explaining
all cases of parapatry by the same n1echanisn1 . One phyletic lin­
eage may evolve very rapidly while others , even closely related
ones , may experience com plete stasis fo r n1any millions of yea rs .
In short, there a re several possible solutions for many evolu tion­
ary challenges , but all of then1 are co mpatible \Vith the Dar\vinian
paradig 111 .
Most evolutionists, pa rticularly those who \vork on a single
group of organis m s , tend to neglect the extraordinary pluralis n1
 Ne w Fron tiers in Ello lutionary f3iolo..<,? y

of evolution . As Franc;ois Jacob ( 1 977) has so rightly said, evolu­

tion is a tinkerer, and in a given situation m akes use of that which
is most readily available . One can take almost any evolutionary
pheno n1enon and show how greatly it differs among the differen t
groups o f anin1 al s and plants. The lesson one learns from this is
that s weeping cl aims are ra rely correct in evolu tionary biology.
Even when s o n1ethin g occurs " usually," this docs not n1ean that
it n1ust occur al ways . One must re member the forever-presen t
pluralism o f evolutionary processes .
A l n1ost every scientific theory is con tinuously in need of revi­
sion and supplen1 entation , yet these chan ges do not necessarily
touch the true core of the theory. The Mendelian theory of inher­
itance is an apt illustration o f this. When it was rediscovered in
1 900, it was expressed in the fo rm of three laws . Wi thin j ust a few
yea rs two o f these laws , those of dominance and independent as­
sortmen t , were found not to be universally valid. Still , no one
claimed that M endel 's theory had therefore been refuted . The
continuing minor revisions and supplementations of the D arwin­
ian theory, including the version given to it during the synthesis,
do not qualify as refutations either.
The architects of the evolutionary synthesis have been accused
by some critics of claiming that they had solved all the remaining
problems o f evolu tion . This accusation is quite absurd ; I do not
know of a single evolutionist who would make such a claim . All
that was claimed b y the supporters of the s yn thesis was that they
had arrived at an elaboration of the Da rwinian paradigm which
seemed to be s u ffi ciently robust not to be en dan gered by the re­
maining puzzles . No one denied that there were many open ques­
tions , but there was the feeling that no matter what answer would
emerge in response to these q ues tions , it would be consistent
wi th the D a rwinian paradi g m . Up to now, it seems to me, this
con fiden ce has not been dis appointed .

The I m pa ct o f M olecul a r B iology on Evol utionary Theory

Early in the history of molecular biolo gy there was a widespread

feeling th at its new discoveries might necessitate a co mplete re-

• 1 49 .
 New Fron tiers in Evolutionary Biology

writing of evolutionary theory. Considering how quickly molec-

.. '
ular biology developed into :i·large and powerful field of its own,
a call for an integration of this new field with classical evolution­
ary biology was to be expected. As White ( I 981) said, " We once
again urgently need a new synthesis of two traditions-those of
evolutionary and molecular biology." Other molecular biologists
went even further and stated that the findings of molecular biol­
ogy had already refuted much of accepted Darwinism. Thus, it
was claimed, " Many of the observations [of molecular biology ]
(inducible mutation systems, rapid genomic changes involving
mobile genetic elements, programmed changes in chromosome
number) challenge the most fundamental assumptions which
evolutionary theories make about the mechanisms of hereditary
variations and the fixation of genetic differences" (Shapiro 198 3 ) .
Discoveries that seem to be in conflict with the picture of clas­
sical genetics are made daily in molecular biology. Perhaps none
was as startling as the discovery that genes are highly complex
systems consisting of exons, introns, and flanking sequences, and
that there are numerous kinds of genes; some have seemingly no
function at all, while several functional classes can be distin­
guished among the active genes. But have any of these discoveries
required a revision of Darwinism? I do not think so.
There is no question that molecular biology has given us nu­
merous great new insights into the working of evolutionary
causes, particularly the production of genetic variation. Gratify­
ingly, in n1ost cases they were a confirmation or elaboration of
existing views. Let me merely mention some of the most impor­
tant molecular discoveries.

( 1 ) The genetic program does not by itself supply the building ma­
terial of new organisms, but only the blueprint for making the
phenoty pe.
(2) The pathway from nucleic acids to proteins is a one-way street.
Proteins and information that they may have acquired are not
translated back into nucleic acids.
(3 ) Not only the genetic code but in fact n1ost basic molecular mech­
anisms are the same in all organisms, from the n1ost primitive
prokaryotes up.

• 150 •
 Ne w Fro n tiers in E110 /utionar y Biology

(4) Many n1utations (changes in the base pairs) seen1 to be neutral or

near-neu tral , that is, without noticeable effect on the selective
value of the genotype, but this varies from gene to gene (sec be­
low) .
( 5 ) A critical co mparative analysis o f 111olecular changes du ring evo­
lu tion provides a very large num ber of pieces of information
suitable for the reconstruction of the phylogeny. This is particu­
larly useful if the morphological evidence is indecisive . How­
ever, molecular characters are also vulnerable to homoplasy­
parallel or convergent production of the same cha racter or phe­
notype.

In teres tingly, u p to the p resent tirne, none of these findings has

required a n essential revision o f the Darwinian paradign1 . More­
over, the relation between rnolecular and evolu tionary biology
has not been entirely one-sided . Darwinian thinking made a ma-
j o r con tribution to the evolution of biochemistry into molecular
biolog y. The study o f the phylogeny o f molecules and the search
for the selective significance of molecular structure have greatly
enriched m olecular biology. N o longer does one encounter in the
literature of m olecular biology any thorough study of a particular
molecule o r group o f molecules that is not at the same time con­
cerned with the evolutionary explanation of the molecular struc­
tures encountered during that analysi s .

NEUTRAL EVO L UTION

I n classical population genetics it was assumed that all mu tations
were either s omewhat favorable and would then be incorpo rated
into the genotype (unless lost by rando m even ts) , or more or less
deleteriou s , and then eliminated whenever becoming ho mozy­
gous . This assumption is, fo r ins tance, reflected in the writings
of R. A . Fisher and H. J. M uller. However, genetic variation
found in nature actually seemed to be larger than could be ac­
counted for by such a p rocess o f relatively rapid elimination o f
variation . This induced D obzhansky t o hypothesize that hetero­
zygotes were so metimes superior (" overdo minant ") to the
homozygotes and that through such " balanced selection " the
holding capacity of the gene pool was considerably in creased .

• 151 •
 New Frontiers in Evolu tionary Biology

In 1 966 when enzyme genes were studied in natural popula­

tions by the new method of electrophoresis (Hubby and Lewon­
tin 1 966 ; Harris 1 966), it was found that variant genes (alleles)
were enormously frequent. For this and other reasons Kimura
( 1 96 8) and King and Jukes ( 1 969) proposed a theory of "neutral
evolution" in which it was claimed that many, perhaps most,
amino acid and nucleotide substitutions in evolution are random
fixations of neutral or nearly neutral mutations. A new allele
(or base-pair substitution) is called neutral if it is functionally
equivalent to the allele (or base pair) it replaces and does not
change the fitness of an organism. Although, this theory was at
first vigorously opposed by most evolutionists, including my­
self, the high frequency of "neutral" base-pair replacements is
now well established. On the other hand, the selective signifi­
cance of numerous alleles that had been considered neutral by
neutrality enthusiasts has also been established (for instance by
Nevo 1 9 8 3 ).
One 1najor reason for the heat of the argument between neu­
tralists and their opponents was that they had a different interpre­
tation of the target of selection. The neutralists are reductionists,
and for then1 the gene-more precisely the base pair-is the tar­
get of selection. Hence, any fixation of a "neutral" base pair is a
case of neutral evolution. For the Darwinian evolutionists, the
individual as a whole is the target of selection, and evolution
takes place only if the properties of the individual change. A re­
placement of neutral genes is considered merely evolutionary
noise and irrelevant for phenotypic evolution. W hichever inter­
pretation one adopts, molecular researches have revealed that the
nun1ber of neutral changes seem to be far greater than the number
of gene changes that have adaptational significance.
The 1nore in1portant a gene or part of a gene is functionally, the
less likely it is that a mutation could occur that would in1prove it.
Nearly all rnutations of such a gene \vould be deleterious and
would be selected against. One potential significance of neutral
evolution about which we are still ignorant is the possibility that
son1e new alleles produced by neutral n1utation n1ay later in evo-

• 152 •
 Neu, Frontiers in Evolutiona ry Biolo(_� y

lution have a positive selective value on a different gcnotyp1c

background.
The Darwinian wonders to what extent it is legitimate to des­
ignate as evolution the changes in gene frequencies caused by
nonselected random fixation. In some of the older (particularly
nineteenth-century) literature on evolution, one finds discussions
on how to discriminate between evolution and n1cre change.
There it was pointed out that the continuing changes in weather
and climate, the sequence of seasons of the year, the geomorpho­
logical changes of an eroding mountain range or a shifting river
bed, and similar changes do not qualify as evolution. Interest­
ingly, the changes in nonselected base pairs and genes are more
like these nonevolutionary changes than they are like evolution.
Perhaps one should not refer to non-Darwinian evolution but
rather to non-Darwinian changes during evolution.

Punctuated Equilibria
Paleontologists, from Darwin's days on, have pointed out that
continuous series of fossils either remain unchanged in time or
show only minor changes in size or proportions. All major evo­
lutionary changes seem to occur rather abruptly, not connected
by intermediates with the preceding fossil series. It was this ob­
servation that induced Schindewolf and several other paleontolo­
gists to propose saltational evolution. How can an adherent of
Darwinian gradual evolution explain the riddle of these gaps?
The theory of peripatric speciation, that is speciation in geo­
graphically isolated populations, permitted Mayr ( 1 954, 1 96 3 ) to
suggest that such inbreeding incipient species are sometimes the
scene of particularly rapid genetic turnover, which would leave
no evidence in the fossil record, owing to the geographical re­
strictedness and short duration of the founder populations.
On this foundation Eldredge and Gould ( 1 972) erected the
theory of punctuated equilibria, stating that most major evolu­
tionary events take place during short bouts of speciation and that
successful new species, after they become widespread and popu-

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 New Frontiers in Evolu tionary Biolog y

lou s , enter a period of stasis, .,I,.a sting sometimes for many millions
of years , during which they show only minimal change . Such
speciational evolution , because it occurs in populations, is grad­
ual in s pite of its rapid rate and therefore is in no conflict whatso­
ever with the D arwinian paradi g m . How often such maj or
changes in founder populations occur and what percentage of
new species enter a subsequent period of stasis are still controver­
sial , however.

Sociobiology
Perhaps the most im portan t develop ment in evolutionary biol­
ogy is the adoption of a largely evolutionary approach in almost
all branches o f biology. For ins tance, the ecologis t asks to what
extent an ecosys tem is the res ult of such evolutionary forces as
com petition and predation ? By what kind of selection pressures
is the partitioning of resources con trolled? Indeed , every ecolog­
ical problen1 is subj ected to a recons truction of the selection fac­
tors seemingly involved, and to a study of their possible adaptive
significance. The same is true for the field of animal behavior.
Not only are the ph ysiological causations of behavior studied but
also the adaptive significance of each kind of behavior, and the
selective a gents that control behavior in relation to other adapta­
tions o f a s pecies .
It is the merit o f Edwa rd 0 . Wilson's magnificent work Socio­
biology : The Netv Synthesis ( 1 97 5 ) to have pointed out that an evo­
lutionary approach is particularl y important in the study of social
behavior. Wilson defined sociobiology " as the systematic study
of the biological basis of all social behavior. " Unfortuna tely, the
slight a mbiguity of the word " b iological" resulted almost imme­
diately in a heated controvers y. For Wilson , " biological " mean t
that a genetic disposition n1akes a contribution to social behavior.
For his politically motivated op ponents , "biological " meant that
social behavior is " genetically detennined . " Of course, we hu­
n1 ans \Vould be mere genetic automata if all of ou r actions were
strictly detern1ined by genes . E veryone agrees that this is not the
case . And yet we also kn ow, particularly from studies of t\vins
sepa rated at birth and adopted into differen t fa n1 ilies , and fro n1

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 New Frontiers in Evolutiona ry Biology

other adoption studies, that a rema rka bly large proportion of ou r

attitudes, qualities, and propensities is affected to some degree by
our genetic heritage. The modern biologist is far too sophisti­
cated to want to revive the old polarized nature-nu rtu re contro­
versy; we know that almost all human traits are influenced both
by inheritance and by the cultu ral environment (Alexander 1 979) .
The most important point made by Wilson is that in n1 any re­
spects the same problems are encountered in human-behavior
studies as in animal studies. Likewise, 1nany of the answers that
seem to explain animal behavior are applicable in the study of
human behavior.
Although the term "sociobiology " was coined only in 1 97 5 ,
the study of the phenomena now often bracketed together under
this term had begun long before; indeed some of these problems
had already been constructively analyzed by Darwin. The num­
ber of problems in this field is legion, but I will mention only a
few. Perhaps the core problem is the question of the evolution of
altruism . Selection by its natu re is a thoroughly selfish process,
measured only in terms of the reproductive advantage it gives to
an individual . The great question, then, is how can true altruism
evolve? An altruistic act has been defined by Trivers ( 1 9 8 5 ) as an
act "that benefits another organism at a cost to the actor, where
cost and benefit a re defined in terms of reproductive success. "
This leads to the question: How could natural selection favor an
act that is detri mental to the actor?
One suggestion was that it could develop by way of so-called
" reciprocal altruism ," by which actor A performs an altruistic act
toward individual B in the expectation that this individual will
reciprocate by performing in turn an altruistic act toward A . To
me, application of the term altruism to such an exchange of fa­
vors is misleading , because such behavior is not detrimental to
the respective actors if there is appropriate reciprocation.
Two other mechanisms have been suggested as explanations for
the evolution of altruism . As far back as 1 9 3 2 J. B . S. Haldane
suggested that altruistic acts toward close relatives might favor
the su rvival and spread of those of the altru ist's genes that are
sha red by these relatives . Since fitness is measured in terms of the
survival of the genes of an individual, it is legitimate to refe r to

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 New Fron tiers in Evolutionar y B iolog y

the successful transfer of one's genes to one's descendants and

other relatives as "inclusiv� htness." The process of contributing
toward the survival of one's close relatives (who share portions of
one's genoty pe) has been referred to as "kin selection."
Parental care, of course, is a special kind of altruism that leads
to an enhancement of reproductive success. There is an extended
literature on the interaction of mates with each other, with their
offspring, and with other relatives, all of it leading to kin selec­
tion. Properly speaking, this is merely part of the enormously
important topic of "selection for reproductive success." There are
factors that favor kin selection, and others that oppose it. The
final outcome is a co1npromise between these conflicting selec­
tion pressures. Maynard Smith's ( 1 9 82) application of game
theory to the subject of altruism and selfish behavior has been
particularly illuminating.
Strictly speaking, kin selection operates only in groups of close
relatives because the genetic advantage of helping another indi­
vidual decreases rapidly with genetic distance. However, there is
another mode by which altruistic behavior can be favored in evo­
lution. In so1ne groups, consisting of both relatives and unrelated
n1embers, certain modes of social facilitation have become estab­
lished that help the entire group. A typical example is the exis­
tence of individuals that serve as "watchdogs" while the rest of
the group is feeding. Such a monitor, when giving warning calls
to alert the other members of his group to a predator, may expose
himself to increased vulnerability, but this altruistic behavior
benefits the survival and reproductive success of his group as a
whole.
Such social "ethics" exist in a rather rudimentary way in many
living organisms, and are described in great detail in Wilson's So­
ciobiology . They are of infinitely greater importance in the human
species, where every cultural group has its code of ethics which,
in the long run, determines the survival and ultimate success of
the group. In the human species, quite clearly, one must distin­
guish two components of ethical behavior, an ancestral one based
on inclusive fitness, particularly parental care, and a cultural one ,
codified in all civilizations in laws and religious dogmas. Un-
 Ne w Fron tiers in Ei,o lutiona ry �iolo<{?Y

doubtedly, a genetic propensity for accepting and n1ainta1n1ng

such cultural prescriptions is favored by selection (Waddington
1 960), but the contents of the ethical repertory are acquired in the
individual's lifetime and are not fixed genetically.
The study of groups exhibiting altruistic behavior has revived
a long controversy about the validity of so-called group selection.
Some authors have postulated that not only individuals but entire
groups may sometin1es serve as the target of selection . We now
understand that this clain1 is valid only for groups with a fitness
value that is greater than the arithmetic mean of the fitness values
of the individuals of which it is composed. There arc two such
groups-those consisting of relatives, where inclusive fitness
contributes to the fitness of the group, and groups of nonrelatives
which practice social facilitation or various forn1s of mutual help.
The controversy as to the validity of group selection has been
resolved by a careful discrimination between these various kinds
of groups. Some species are, while others are not, subject to
group selection.

Questions a t the Frontier

If someone were to ask me which frontiers of evolutionary biol­
ogy are likely to see the greatest advances in the next ten or
twenty years, I would say elucidation of the structure of the gen­
otype and of the role of development. The simplistic reductionist
view of the relation among genotype, development, and evolu­
tion is that each gene is translated into a corresponding compo­
nent of the phenotype and that the contributions of these com­
ponents to the fitness of the resulting organisn1 determine the
selective value of the genes. There is undoubtedly some truth in
this view, but it greatly oversimplifies the actual connections.
This had been realized all the way back to l)arwin's day. Darwin
himself spoke of the mysterious laws of correlation, and Haeckel
converted the Meckel-Serres Law into the evolutionary principle
of recapitulation. Again, this principle was a vast oversin1plifica­
tion, but it has a correct nucleus. The fact that the land-living
vertebrates go through a gill arch stage in their developn1ent is a

• I 57 .
 New Frontiers in Evolutionary B iology

powerful clue for their descent

.. '
from aquatic ancestors, to give
only one example. Both ofthese generalizations call attention to
holistic interactions. B y contrast , the belief of some reductionists
that the role of genes is exhaustively described by the largely in­
dependent contribution of each gene to some particular aspect of
the phenotype neglects the fact that the genotype is a complexly
. .
interacting system .
Each year one or two books or sy mposia volumes are pub­
lished on the relation between genotype, development, and evo­
lution, and it is quite impossible in this short chapter even to ou t­
line the basic problems. To make matters worse, though the
questions are increasingly better understood, no satisfying an­
swers have yet been found for most of them . When answers have
been suggested , they often contradict one another. Instead of at­
tempting to present a detailed account of the problems and the
opposing attempts to solve them, I will satisfy myself by merely
indicating some of the general directions of future research.

DOMAINS OF THE GENOTY PE

We know now that there are different classes of genes, and they
play not only different roles in ontogeny but also in evolution.
Fu rthermore , certain genes seem to be tied together into func­
tional units and seem to control development as units. They seem
to represent well-circu mscribed domains that lend a hierarchical
structure to the genotype . The existence of such domains is not
necessarily in conflict with Mendelian segregation. How the de­
velopn1ental and evolutionary conservation of such domains is
effected is not yet understood, though the discovery of the wide­
spread occurrence (from yeasts to mammals) of homeoboxes
(Robertson 1 98 5 ) and the study of complete sets of immune genes
reveal possibilities.
At the present tin1e the existence of such domains is primarily
indicated by indirect evidence . Among a number of categories of
such evidence, I would like to discuss in son1e detail one particu­
lar phenon1enon, called mosaic evolution.
For a physicist, one of the n1ost important paran1eters of any
process is its rate. In the endeavor to n1ake evolutionary biology
as si111 ilar to the physical sciences as possible, evolutionists have

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 New Frontiers in Evo lutionary B_[ oloJ!Y

attempted fro m 1 8 5 9 on to determine evol ution ary rates ( S imp­

son 1 944 , 1 9 5 3 ) . Alas, this has been a rather frustrating ex peri­
ence . Rates of evolution , speciation, and extin ction may differ by
several orders of n1agnitude in differen t organis n1s and under dif­
ferent circumstances . Perhaps most distu rbing was the discovery
that organisms do not evolve as harmonious wholes but that dif­
ferent cha racters (con1ponents o f the phenotype) and domains of
the genotype may evolve at highly different rates . This was
already know n to Lamarck and Darwin and has since been con­
firmed by nu merous evolu tionists, particularly the paleontolo­
gis t s . A rchaeop tery x is an apt illustration . In some of its character­
istics (feathers a n d wings) it is already a typical bird, in others
(teeth a n d tail) it is still a reptile, and in still other characters it is
more o r less intermediate between the two classes . A . C . Wilson
( 1 974) has pointed out that frogs are mo rphologically highly con­
servati ve, and this is also true fo r their ch romoso mes , but that
different phyletic lines of frogs seem to diverge in their enzy me
genes a t about the same rate of divergence as mammals , which
evolve very rapidly in all three types of characters . What this sug­
gests is that different do1nains of the genotype seen1 to evolve
independently to a considerable degree. This is well illustrated ,
for instance, in a co mparison of man with his anthropoid rela­
tives . O ver a l l , man is most similar to the chimpan zee , but in a
few chara cters man's genotype is more similar to that of the go­
rilla or even the orangutan .
The term " mosaic evolution " has been used for the indepen­
dent evolution of different domains of the genotype . At the mo­
lecular level it was found not only that differen t molecules have
different evolutionary rates but also that the same molecule may
change its rate at d i fferent stages in the evolution of a phyletic
lin e . Whenever there is a period o f maj or adaptive radiation , it is
acco m panied by a highly accelerated rate of molecular evolution ,
later followed b y a deceleration to slow change . The so-called
molecular clock m ay keep very different times for differen t n1ol­
ecules and may change its rate within a phyletic line . As Good­
man ( 1 9 8 2 : 3 77) has correctly poin ted out, none of these findings
on chan ges in molecular timing invalidates the l)arwinian thesis
that natu ral selection is the princi pal force directing the evolution

. 159 •
 New Fron tiers in Evolutionary Biology

of n1olecules and organis m s . Mos aic evolution , however, must

be carefully considered in the'construction of phylogenetic trees .
The o pposite of the independent evolution of domains of the
genotype is the integration of these do mains and the seen1ing
cohesion of the genes that belong to any one of these domain s .
Just exactly what controls this cohesion i s s till largely unknown,
but its existence is abundantly documented . The almost total in­
tegri ty of the gene co m plex that controls the nu mber of extren1-
ities in tetrapods , insects , and arachn ids is one of litera lly hun­
dreds of exa mples . During the pre-Cam brian period , \vhen the
cohesion of the eukaryote genotype was still very loose, seven ty
or n1ore distinct morphological ty pes (phyla) fo nned . Through­
out evolution there has been a tendency for a progressive B con­
gealing " of the genotype, so that devia tion fron1 a long­
establis hed morphological type has becon1e more and n1ore
difficult . This is one of the well-known constraints in evolu tion­
ary change, and one of the reasons why natural selection has such
lin1ited leeway. When we speak of the congealing of the genotype
or, i f we prefer, of the progressive in tegration of the genotype,
we are simpl y using words to cover up our ignorance . Neverthe­
less, the evidence indicates that the older a taxon is , the n1ore
difficult it is to escape from the straitj acket of a highly in teg rated
(congealed) genotype. This is why not a single new rnorpholog­
ical type (phylum) has originated since the Can1brian , over 5 00
rnillion years ago .
A t first sight this conclusion would seen1 to be contradicted by
the ada ptive radiations that have occurred in so many phyletic
lines . A fter a stasis of I oo million years or longer a higher taxon
rn ay suddenly experience a burst into nun1erous new ta x a , as
happened to the man1 n1als in the Paleocene and Eocene. Hovv­
ever, most of these were ad hoc adaptations and did not really
deviate very far fro111 the ancestral Ba 1 1pla 11 or structu ral type. So
far as I can j udge, the s a n1e is true for n1 ost of the other \vell­
kno\vn radiations . The songbirds , fo r ins tance, have produced
over 5 , 000 living s pecies since their origin , but excep t fo r their
plu rn a ge and bills, they exhibit hardly any devia tion fro111 the
standard Ba11p la11 .

• 1 60 .
 Ne w Fron tiers in E110 /1aiona ry B,iolo(_e y

The power of these constraints is evident at every hierarchical

level, son1eti111es even at the species level, as docun1cnted by the
frequency of sibling species (which are reproductively isolated
but morphologically identical) . Indeed, n1ost geographic varia­
tion deals n1crely with minor quantitative changes in characters,
without affecting the genotype appreciably. Any restructuring ,
when it occurs, apparently takes place aln1ost always in periph­
erally isolated founder populations.
Evidence for increasing or decreasing cohesion of the genotype
can also be found in the study of n1acroevolution. Evolutionary
acceleration or deceleration n1ay be displayed by all or at least
most me1nbers of a new higher taxon. A well-known textbook
L.

case is evolution an1ong the lungfishes, as worked out by Westoll

( 1949). Almost all the anatomical reconstruction in this new class
of fishes took place in the earliest stages (during son1e 25 million
years), while almost no change has taken place in the subsequent
200 million years. Such a drastic difference between the rates of
evolutionary change in young and n1ature higher taxa is virtually
the rule. Bats diverged fron1 insectivores within a few million
years, but their morphology has hardly changed in the last 40
million years. The fossil record (though inadequate) suggests that
the same is true for the anaton1ical reconstruction during the ori­
gin of birds and whales. Exceptions are much in the minority.

THE R O L E OF S O M AT I C P R O G R A MS IN EVO L UTION

The traditional formula according to which developn1ent is pro­
grammed by the genotype implies far too direct a pathway from
the gene to the endpoint of its action in the phenotype. Embryol­
ogists since Kleinenberg ( 1 8 86 ) have known that genes might ef­
fect the production of an embryonic structure which subse­
quently serves as part of the program for the later stages of
development. That not all developmental progran1s are purely
genetic, and that there are also somatic progran1s, is best made
clear by the example of programs controlling behavior. W hen,
for exampie, a male bird performs a certain courtship display, this
action is not programmed directly by the genotype but rather by
a secondary program laid down in the central nervous systen1

• 1 61 •
 New Fron tiers in Evolu tionary Biology

during ontogeny. And it is .t,his secondary-somatic-progra m

that actually controls the behavior.
That somatic programs exist is probably not of major conse­
quence for the evolution of behavior. The existence of somatic
programs, however, may be of high significance in morphologi­
cal evolution . I t may help to explain many puzzling phenomena
of both ontogeny and evolution . For instance , it may explain
most ins tances of recapitulation . When an embryonic structure of
a species ' ancestors is maintained in ontogeny even though it no
longer seems to be of functional value (for ins tance, the gill a rches
of the mam malian embryos) , such an embryonic stru cture may
have been preserved by natu ral selection because it serves as the
somatic program for the subsequent ontogenetic stages . The ex­
isten ce of somatic progra ms imposes important constraints on
evolutio n . Much developmen t in the higher taxa seems to be con­
strained by such son1atic programs , which seem to be highly re­
sistant to evolutionary change . This statement is , of cou rse, at the
present tin1e onl y words . However, recent research in molecular
ontogeny has made promising beginnings . Progress is bound to
be slow, because develop ment involves highly complex interac­
tions between different domains of the genotype and different so-
1nat1c p rogra m s .
This subj ect area i s of particular interes t t o the student of ma­
croevolution . It is here that the connection will be made between
the genetics of individuals and populations on the one hand and
the maj or ma croevolutionary processes and events on the other.
Resea rch will have to pursue simultaneously reductionis t ap­
proaches (that is , the study of the action of individual genes) and
also holistic ones (tha t is , the study of domains of the genotype
and of whole so matic programs) .

Da rwinism Toda y
The basic theory of evolu tion has been confirn1ed so co mpletely
th at 1nodern biologists consider evolution sin1ply a fact . Ho\v
else except by the wo rd " evolution '' can we designate the se­
quence of fa unas and floras in precisely dated geological strata?
And evolutionary change is also sirnply a fact O\ving to the

• 1 62 .
 Neu, Frontiers in Evolutionary ]Jiolo�<.? Y

ch an ges i n the con ten t o f gene p ools fron1 generation to gene ra­
tio n . I t is as m u ch of a fa ct as the obs erva tion that the ea rth re­
vo l ves a r o u n d the s u n rather than the reverse . Evolu tion is the
fa ctual b a s i s on which the other fo u r Da rwinian theo ries res t . Fo r
instance , a l l the p hen o m en a explained b y co m m on des cen t
wo u l d n1 ake no sen se if evo l ution we re not a fa c t .
D a r w in's s e c o n d theo ry-co m m on des cen t-al so h a s been
g l o rio u s l y con fi r m ed by all resea rches since 1 8 5 9 . Eve rythin g we
have lea rned a b o u t the p h ysiology and ch em istry of organ i s m s
su p po rts D a r w in's d a ri n g s pecu lation t h a t " a ll t h e organ ic beings
which have ever l i ved on this earth have descen ded from some
one p r i m o rd i a l fo r m , into which l i fe was firs t breathed "
( 1 8 5 9 : 4 84) . T h e dis covery that the proka ryo tes h ave the san1 e ge­
netic code as the higher o rg a ni s m s was the most dec i s i ve con fir­
m a tion o f D a r win's h y pot hesi s . A histo ri ca l unity in the en tire
l i v i n g wo rld cann o t help but h ave a deep mean i n g fo r any think­
ing person a n d fo r his feelin g towa rd fel l ow o rganis m s .
D a rwin's g reat e m p h a s i s o n the devel o p ment of d i versity a s an
i m p o rtant co m po nen t o f the evolu tion a ry p rocess - his third
theo r y - was undes erved l y ne g lected du rin g t he fi rs t third of this
cen tury but is a g ain at the fo refron t o f in teres t, parti cularly in
paleontol o g y and ecolog y. A s fa r as s pecia tion is concerned -the
p roces s that ser ves as the sou rce o f new diversity - D a rwin wa s
so m e w h a t con fu se d , h aving s u p p o rted geogra phic s p eciation on
islands and a wi des p read s y m pa t ric s peciation on con tinen ts . The
cont roversy over specia tion is s till alive to day, but the basic
theory that s pecies m u l t i p l y as wel l as e vo l ve is uncon tested .
Da r w i n's theory o f g ra d u a lis m , u n p a latable even to his close
friends H u x l e y and Galton , has u l t i m a tel y triu m phed decisive l y
and m a kes m o re sense the m o re clea rly w e realize that evo l u tion
is a p ro ces s invo l ving p o p u l a t i on s . The onl y a p p a ren t exceptions
a re the o ccasional a b an d on m en t of sexual reproduction and cer­
tain chro m o s o m a l p rocesses such a s p o l y p loidy. B u t th ese p ro­
cesses have not l e d to any m a croevolut ion ary conseq uences d i f­
fe ren t fro m p o p u l a tional evo l u tion . A l l other s pecia tion is
p o p u l a ti on a l , even in the theory o f pu nctua ted equilibri a .
The g reatest t ri u m p h o f D a r winism is that the theory of na tu­
ral select ion , fo r eigh ty yea rs a fter 1 8 5 9 a minority o p inion , is
 New Frontiers in Er,10 /utionary Biology

now the prevailing explanation of evolutionary change. It has

achieved this position both• kiy irrefutable proofs and by default,
as all the opposing theories were demolished. Darwin took it for
granted that a nearly unlimited amount of variation was at all
tin1es available to provide raw material for natural selection. He
had no idea as to the source of this variation and supported several
genetic theories (soft inheritance, pangenesis, blending inheri­
tance) that have since been refuted. Nevertheless, advances in ge­
netics continue to strengthen rather than weaken the theory of
natural selection. Darwin was ren1arkably astute in his concep­
tualization of selection. He clearly saw (better than A. R. Wallace
and most other contemporaries) that there are two kinds of selec­
tion, one for general viability leading to survival and the rnainte­
nance or in1provcn1cnt of adaptedness, and this he called "natural
selection," and another that leads to greater reproductive success,
and this he called "sexual selection."
Where evolutionists today differ fro111 l)arwin is altnost en­
tirely on rnatters of ernphasis. While Darwin was fully aware of
the probabilistic nature of selection, the 1nodern evolutionist en1-
phasizes this even n1ore. The n1odcrn evolutionist realizes how
great a role chance plays in evolution. Darwin never said, "Selec­
tion can do anything" ; neither do we. On the contrary, there are
powerful constraints on selection. And selection is, for various
reasons, appallingly often unable to prevent extinction.
One hundred and thirty years of unsuccessful refutations have
resulted in an in11nense strengthening of Darwinisn1. Controver­
sies within evolutionary biology about such n1atters as the occur­
rence of syn1patric speciation, the existence or not of cohesive
don1ains within the genotype, the relative frequency of con1plete
stasis in species, the rate of speciation, the significance of neutral
allele replace111ent, and \vhatever else all take place within the
fra1nework of Darwinis111. The basic Darwinian principles are
n1orc firn1ly established than ever.

• 1 64 •
 References

Glossary

Acknowledgn1ents

Index
 References

Adan1s, M. 1 9 8 0. Sergei Chetverikov , the Koltsov Institute, and the evolu­

tionary synthesis . I n E. Mayr and W. B. Provine, The Ei1 0/u tionary
Syn th esis, pp. 242-2 7 8. Cambridge : Harvard University Press.
Alexander, R. D. 1 979. Da rw in ism and Human AJfairs . Seattle : University of
Washin gton Press.
Ayala, F. J. 1 9 8 2. Beyond Darwinism ? The challenge of macroevolution to
the synthetic theory of evolution. Ph ilosophy (f Science Association
(PSA ) 2 : 2 7 5-29 1 .
--- 1 9 8 3 . Microevolution and n1acroevolution. I n D . S. Bendall, ed. ,
Evo lu tion from Mo lecu les to Men , pp. 3 8 7-402. Cambridge : Cam bridge
University Press.
Barlow, N. , ed. 1 96 3 . Darwin's ornithological notes. Bull . Brit . Mus . (Nat.
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Westoll , S . 1 949 . O n the evolutio.Q of the Dipnoi. In G . Jepsen , E . May r,

and G . G. Simpson , eds . , Genetics , Paleontology, and Evolu tion , pp. 1 2 1 -
1 84 . Princeton , NJ : Princeton University Press .
Whewell, W. 1 840. Ph ilosophy of th e Inductive Sciences . London : Parker.
White, M . J. D . 1 97 8 . Modes of Speciation . San Francisco : Freeman.
--- 1 98 1 . Tales of long ago . Pa leobiology 7 : 2 8 7-29 1 .
Willis , J . C . 1 940. The Course of Evolution by Differentiation or Divergent Evo­
lution Rather Than by Selection . Cambridge: Cambridge University
Pres s .
Willmann, R . 1 9 8 5 . Die A rt i n Raum und Zeit. Berlin und Hamburg : Parey.
Wilson, A . C . , V. M . Sarich, and L . R . Maxson . 1 974 . The importance of
gene rearrangemen t in evolu tion : evidence from studies on rates of
chromosomal, protein , and anatomical evolution . Proc . Nat . A cad . Sci .
7 1 : 3 02 8 -3 03 0 .
Wilson, E . 0 . 1 97 5 . Sociobiology : Th e New Synthesis . Cambridge, MA :
Harvard University Press .
Young , W. 1 9 8 5 . Fallacies o_f Creation ism . Calgary, Alberta : petselig Enter­
prises Ltd.
 Glossar y

Acquired characters Those characteris tics of an organis m 's appea ran ce

(phenoty pe) that resu l t fro m environ mental infl uences rather than in heri­
tance

Adaptationist program The resea rch en deavo r to discover the ada ptive
signifi cance o f s t ru ctu res and behav iors

Adaptedness The suitability of a s tructu re or an organis m fo r its envi­

ron ment or l i fest yle, as a resu l t of past selection

Additive inheritance S t ress on the independen t effect of individual

genes , with disregard fo r the m u tu a l ( epistatic) in teraction o f genes

Allele A ny of the alternative v a riants of a gene

Allopatric speciation See Geographica l s peciation

Altruism Behavior that benefi ts another organis m at a cost to the actor,

where cost and benefi t a re defined in terms of reprodu cti ve success

Artificial selection Selection o f b reed ing stock by an animal or plant

breeder

Base pair A pair of h yd rogen-bonded nitrogenous bases ( one purine and

one p y ri m idine) that connect the t wo strands of the DNA double helix . In
D N A , aden ine pairs only w i th t h y m ine, and guanine pairs only with cy­
tosine. This co m plementarity is the key to the sel f-replica tion and infor­
m a tion-trans m i t ting capabilities o f D N A

Bauplan The basic s t ructural characteristics ( m o rphot ype) o f a given

t y pe of organis m , such as the Bau p lan of the verteb ra tes , or of the bi rds , o r
of t h e insects

Biological species concept Defines a species as a rep roductively iso­

la ted a g g regate of populations which can interbreed beca use they share the

. 1 77 .
 Glossary

same isolating mechani s m s . See E.yolutionary species concept; Nominalist

species concept; Typological species concept
Biota Fauna and flora
Blending inheritance A now-disproven theory that the genetic deter­
m inan ts o f the parents fuse into a uniform substance during the fertilization
of the egg . See Particulate inheritance
Catastro p hism A theory that catastrophic events in the history of the
earth have resulted in the partial or complete extinction of the biota
Causations, p roximate See Proximate causations; Ulti mate causations
Central dogma The assertion that the info rmation contained in proteins
cannot be translated into nucleic acids
Character A com ponent of the phenotype
Character divergence The name given by Darwin to the differences de­
veloping in two (or more) related species in their area of coexistence, owing
to the selective effects of competition
Chromosome One of the threadlike structures in the nu cleu s of the cell,
consisting of DNA and associated protein . See DNA; Gene
Common descent The derivation of certain species or higher taxa from
a com mon ancestor
Cosmic teleology The belief that the universe as a whole or some of its
changes are directed towa rd a final objective or goal , such as greater perfec­
tion
Creationism A belief in the literal truth of the story of creation as re­
corded in the Book of Genesis
Deoxyribonucleic acid See D N A
Deism A belief i n a supreme being who rules the world th rough divinely
ordained laws rather than through direct in tervention
Determinism A theory that the ou tco me of any process is strictl y pre­
detennincd by definite causes and natural la\vs and is therefore predictable
Di p loid Having a double set of ch ro mosornes . See Haploid
DNA (deoxyribonucleic acid) The rn olecule that ca rries the genetic
inforrna tion (genes) in all organisn1s except the RN A viruses . It con sists of
t\vo long pol ysugar-phosphate strands connected by base pairs and tw isted
in a double helix. See Chro n1oso1ne; Gene

• l /.. l� •
 G lossa r y

Epigenesis Develop1nent from an un formed basic 111a terial, as opposed

to preformation . See Prefo rmation
Essentialism The belief, going back to Plato , that the ch angin g variety
of nature can be sorted into a limited number of classes , each of which can
be defined by its essence . Varia tion is simply the mani festa tion of in1 perfect
rep resen tation of these con stant essences . Also referred to as ty pological
thinking
Eukaryotes O rganisms whose cells have a nucleu s with discrete ch ro­
n1osomes as well as such cellular organelles as mi toch rondria and chloro­
plasts
Evolutionary causations See Ulti mate causations
Evolutionary species concept Defines a species as a ph yletic lineage
evolving separately fro m others , with its own evolution ary tenden cies and
historical fate. See Biological species concept; Non1inalistic species con­
cep t ; Typolog ical species concept
Evolutionary synthesis A somewhat modified Da rwinian paradig m
which includes a refu ta tion of transformational evolution, saltation is m , and
orthogenesi s , while placin g g reat emphasis on natural selection , adapta­
tion, and the study of diversi ty (origin of species and h igher taxa) . This
paradigm was worked out in the 1 930s and 1 940s by a g roup of " a rchitects "
of the evolutionary syn thesis, which included Dobzhan sky, Mayr, Rensch ,
Simpson , S tebbins, and Ti mofeeff-Ressovsky. The term is sometimes also
used to refer to the ti me period when the paradigm was being constructed,
an d the paradi g m is so metimes referred to as the modern synthesis
Exons Sequences of base pairs in a gene that participate in the coding o f
the peptides . See In trons
Female choice A theory in sexual selection which states that it is often
the female that selects one of several available males for mating , rather than
the other way around
Finalism A belief in an in heren t trend in the natural world towa rd some
preordained final goal o r purpose, such as the attain men t of perfection . See
Teleology
Fisherism The set of evolution ary theories held or proposed by R. A .
Fisher wh ich stressed the power o f selection , the gene as the unit o f selec­
tion , the particulateness of inheritance, and the aln1ost exclusive p revalence
of additi ve inheri tal}ce . Evolu tion fo r Fisher was a change in gene frequen­
cies within a popul ation

• 1 79 .
 Glossa ry

Fitness The relati ve ability of •a'n

o rganism to survive and transmit its
genes to the gene pool o f the next generation
Gamete A germ cell (egg o r sperm) carrying half of the organ ism's fu ll
set of chromosomes; es pecially a mature germ cell capable of participating
in fertilization . See Genetic recombination ; Meiosis
Gene In classical genetics , a unit of inheritance, transmitted from gener­
ation to generation b y an ovu m or sperm, which controls some cha racter­
istic of an individual or some aspect of the individual 's development. In
molecular biology, a sequence of base pairs in a DNA molecule that con­
tains information for the construction of one protein molecu le (peptide) .
See Base pair; Chro mosome; D N A
Gene pool The totali ty of the genes of a population or a species
Genetic code The code that determines which triplets of base pairs are
translated into which a n1ino acids
Genetic drift Changes in the gene content of a population owing to
" chance , " that is , to stochastic processes
Genetic program The information coded in an organis m 's D N A
Genetic recombination The reshuffiing o f a n organis m 's genes during
the production o f germ cells, through crossing over of sections of the or­
ganis m 's ma ternal and paternal chromosomes . It occurs during meiosis ,
just prior to cell division and the chromosomes ' independent assort ment
during the reduction division . Genetic reco mbination assures that the chro­
mosomes carried by an organis m 's eggs or sperm are not identical to the
chromoso mes the o rganism inherited from either of its paren ts . No two
ch romoso mes of any of the eggs or sperm are likely to be identical to one
another. Genetic reco m bination , as now recognized , is res ponsible for a
large share of the variation upon which natu ral selection acts . See Meiosis
Genome The totality of genes carried by a single gamete
Genotype The genetic cons titu tion of an individual, especially as distin­
guished from its ph ysical appearance. See Phenotype
Geographical speciation Speciation that occurs while populations a re
geographically isolated; also known as allopatric speciation
Germ cell A cell whose principal function is reproduction ; an egg or
sperm cell
Germ plasm An outmoded term \vhich referred to the genetic material
in the gern1 cells . as distin guis hed fro m the " so rna," that is, the phenotype

• 1 80 •
 Glossa ry

Gradualism A theory that evolution progresses by the gradual modifi­

cation of populations, and not b y the su dden o rigin of new ty pes (sal ta­
tions) . See Saltational evolution
Great chain of being See Scala natu rae
Ha p loid Having the nu mber of chro m osomes of a norn1al genn cel l ,
which equals h a l f the n u m ber i n a somatic cell
Hard inheritance A theory that the genetic 1n aterial is con stant ( " hard ")
and cannot be a ffected by lifestyle or the environmen t . None of the chan ges
in the phenotype of an organisn1 during its lifes pan can be passed on to its
offspring . In the terminology o f molecular biology, hard inheritance is the
theory that informa tion in the proteins cannot be conveyed to the nucleic
acids in D N A ; also known as the " central dogma . " See Blending inheri­
tan ce; Inheritance of acquired characters ; Particu late inheritance; Soft inher­
itance
Heteroz y gosit y The occurren ce of two differen t vanants (alleles) of a
gene at the corresponding loci on two ho mologous chron1oso mes . See
H omozygosity
Homeobox A defini te seq uence of genes con trolling a step in develop­
ment, particularly in meta meric organisms
Homoz y gosit y The occurrence of two iden tical alleles at the corre­
sponding loci on two ho mologous chron1 oso 1nes . See Heterozygosity
Horizontal evolution The simultaneous evolution of geographically
dis tributed populations; geo g raphic variation
H y brid The p roduct of the interbreeding of different kinds o f organis 1ns,
ordinaril y of different species
Inci p ient s p ecies A popula tion in the process of evolving into a separate
species
Inclusive fi tness The su m o f an individual 's own fitness plus all its influ­
ence on fi tness in its relatives o ther than direct descendants
Infusorians One-celled organis m s ; protists
Inheritance of acq uired characters A now-disproven theory that
changes in an o rganis m 's phenotype which result from factors in its envi­
ron ment can be passed on to o ffspring th rough the organis m 's genetic ma­
terial
Introns N oncoding seq uences of base pairs that are eliminated prior to
the translation of the nucleic acids into proteins (peptides)

• 1 81 •
 Glossary

Isolating mechanisms B iolog-i�"ai or behavioral p roperties of ind ividuals

wh ich preven t the in terb reedin g o f populations that coexist in the san1e area

Kin selection Selection fo r the sh ared co m ponents of the genotype in

individuals related by com mon descen t

Lamarckism A belief in the gradu al change of species ove r time to a

" higher" (meaning better adapted and more co mplex) level through the
inheritance o f acqu i red cha racters

Macroevolution Evolu tion above the species level; the evolut ion o f
higher taxa a n d t h e p rodu ction o f evolutionary novelties such a s ne\v st ruc­
tures . See M i croevo l u tion

Meckel-Serres Law. A law which s tates that there are paral lels bet \veen
the stages in ontogeny and the phy lo genetic series

Meiosis A special series of cell divi sions during the produ ction of the
ga metes in which the n u 1n ber of chron1oson1es is reduced . This process
precedes the produ ction of gan1etes in animals and spores in plants . It nor­
mally involves cros sing over and an independen t assortment of ho1nolo­
gous chro moso1nes . See Genetic recombination

Mendelism In genetics . a stress on hard in heritan ce and the particulate­

ness of the units o f inheritance, as concluded from Mendel 's la ws . The early
Mendelians also believed in drastic mutations (saltational evol ution) and
they minimized the i m portance of natural selection

Metameric C onsisting of a seria tion of parts

Microevolution Evolution at and below the species level . See Macroe­

volu tion

Monotypic U sed to describe a taxon that contains onl y one taxon of the
nex t lower catego ry, such as a genus that co ntains only one species . See
Pol ytypic

Morphology The science of forn1 and structu re in animals and plan ts

Mosaic evolution Evol u tion which proceeds at different rates fo r differ­

ent structu res or fo r other con1 ponents of the phenot ype and genotype

Mutation In mo iecular biolo g y. a change in the geno type . I f the change

occurs in the D N A o f a sornatic cell , the n1 utation may cause a change in
the organisrn 's phenoty pe (leading . for exa 111 ple. to cancer) but will not
affect the o rganis m 's o ffspring; on ly nnl ta tions in the genn cel ls can ca use
heritable chan ges in the o ffsprin g . For the Mendelian s . n1 u tat1on \Vas
thought to be the proces s that p roduces new species

• 1 82 •
 Glossa r y

Natural theology The study of na ture to docu n1en t evidence for the
power and wisdon1 of the C reator in the design of His world
Natural selection The non rando n1 su rvival and reproductive su ccess of
a s n1all percen tage of the individuals of a popu lation owing to their posses­
sion of, at that 1n oment, characters which en hance thei r ability to survive
and rep rod uce . See A rtificial selection ; Sexual selection
Neo-Darwinism A theory of evolu tion develo ped by August Weis111ann
in the late nineteenth century wh ich con sisted of Darwinis m withou t the
inheritance of acquired characters and with a strong e1n phasis on natural
selection
Neutral evolution The occu rrence and accumu lation of heritable n1 uta­
tions which do not affect the fi tness of the individual or its offsp ri ng
N ominalistic species concept Defines a species as the arbitrary brack­
eting of individuals un der a species name. See Biological species con cept ;
Evolutionary species concept; Typological species concept
Niche The constellation of environmen tal factors into which a species fits
or \vhich it requires fo r its survival and rep roductive su ccess
Nondimensional Without a geograph ical or ti1ne dimension . A nondi­
mensional species is a species as encountered at a particular locality and
time, without reference to its geograph ical relation to other populations or
to its evolutionary his tory
Ontogeny The develop men t of the individual from the fertilized eg g
(zygote) to adulth ood
Orthogenesis The evolution of phyletic lineages along a predetermined
linear path way, not through natural selection
Paleontology The study of fossils and ancient life fo rms
Pangenesis A theory adop ted by Charles Darwin to provide a mecha­
nism b y which an inheritance o f acquired cha racters n1ight be explained . I t
invol ved the t ransfer of g ranules (gem 1nules) fro m the b o d y to the gonads
and germ cells
Parapatric speciation The p ro g ressive divergence of two neighboring
popu lations, while meeting in a con tact zone, un til they have beco me two
differen t species
Particulate inheritance A theory developed in the late nineteen th cen­
tury an d adopted by the Mendelians that inheritance is a ffected by genetic
units which do not fuse or blend in the offspring but re main discrete . See
Blen ding inheritance; Hard inheritance

. 1 83 .
 Glossa ry

Peri p atric s p eciation Specia.tJon by budding ; that is, the origin of new
species through the modification of peripherally isolated founder popula­
tions
Phenon ( p l . p hena) A sa m ple of phenotypically similar individuals; a
subdivision of a population or species characterized by phenotypic similar­
ity
Phenot y p e The totality of the observable characteristics of an individual ,
resulting fro m interactions between the environ ment an organism encoun­
ters and the genotype it inheri ted . See Genotype
Ph y logen y The origin and su bsequent evolu tion of the higher taxa; the
history of the evolutionary lineages
Ph y sicalism A philosophy of science widely held in the seventeenth cen­
tu ry and later wh ich asserted that all natural p rocesses (including those in
living organis ms) can be completely and sufficiently described in chemi­
cophysical terms and that any valid statemen t in any field of science can , in
principle, be reduced to an empirically verifiable physical statemen t . Phys­
icalism originally included an acceptance of essentialism, determinis m , and
a reliance on universal laws, and it included no reference to time (historical
processes) . This theory has been largely abandoned , and in other parts con­
siderably modified, owing to develop ments in biology and n1odern physics
Ph y sicotheolog y See Natural theology
Pol y morphism The coexistence of several well-defined distinct pheno­
ty pes or alleles in a population
Pol y p loid y An increase, usually doubling, of the number of chromo­
somes over what is normally found in the somatic cells, due to a doubling
of the chromoso mes of the nucleus, not followed by a division of the cell
Pol y t y p ic Used to describe a taxon that contains more than one ta xon
of the next lower category, such as a genus that con tains several species .
See Monotypic
Po p ulation thinking A viewpoint which emphasizes the uniqueness of
every individual in populations of a sexually reproducing species and there­
fo re the real variability of populations; the opposite of typological thinkin g .
See Essentialism
Po p ulation In evolu tionary biolog y, the community of potentially inter­
breeding individuals, particularly at a given locality
Preformation Development of the em bryo fro n1 n1aterial in \vhich the

. 1 84 .
 Glossa ry

even tual fo r m of the adu l t is " preforn1ed , " that is, already exists in its es­
sen tial structures

Prokaryotes Organis ms without a structu red cell nucleu s , such as var­

ious kinds of bacteria and the so-called blue-g reen algae

Punctuated equilibria See Punctuationism

Punct uationism The theory that most evolu tionarily i m portant even ts
take place d u ring short bou ts of specia tion , and that once species are
fo rmed they are relatively s table, sometimes for very long periods . A lso
known as speciational evolu tion

Quinarianism A n obsolete theory of classifica tion acco rding to which

each ca teg ory con tains a ci rcle of five taxa. Each taxon is in contact ( " os­
cula tes ") with a taxon in a different ci rcle, owing to affinity with it

Recapitulation The passing of an individual o rganisrn , d u ring its devel­

op men t (ontogeny) , th rou gh the s a me s tages of develop men t as did its
ances tors during p h y logeny. See Onto gen y ; Phylogeny

Recombination See Genetic reco mbination

Rectilinear series A series o f fossils in a ph y letic lineage, seeming} y pro­

gressin g in a l inea r m an ner

Reductionism A philosophy wh ich states that all phenomena and laws

relating to co m plex systems (includin g living ones) can be reduced withou t
res idue to those o f the physical s cien ces , and more particula rly, to the
s mallest co mponen ts . See P h ysicalism

Saltation I n evolu tion ary theory, the assertion that new ty pes of orga­
nisms o riginate by the sudden o rigin o f a sin gle new individual wh ich be­
co mes the p rogen itor o f this new kind of organis m

Sal tational evolution Change owing to the sudden ong1n of a new

type, that is, the production of a new kind of individual who gives rise to
a new group o f organi s m s . S ee Transformational evolution ; Va riation al
evolution

Scala na turae A linea r arrangemen t of the fo rms of life from the lowes t ,
nearly inan i m a te, t o the m o s t perfect

Selectionism The theory th a t adaptive chan ges in evol ution are the re­
sult of natural selection

Sexual selection The increased rep roductive su ccess o f a s m all percen t­

age of the individuals of a popul ation owing to thei r possession of, at that
mo men t, cha racters wh ich en hance either their abili ty to co mpete with
 Glossary

. . attractiveness to members of the opposite

members o f the same sex or their�
sex . See Natural selection
Sibling species Morphologically similar or identical populations that are
rep roductively isolated
Sociobiolog y The syste matic study of the b iological basis of all social
behavior
Soma The body ; the phenotype, in contrast with the genotype
Somatic program A stru cture or s tage in development providing infor­
m ation for ensuing development or o ther activities
Speciation The process whereby species multiply; the acqu isition of re­
productive isola tion between populations
Speciational evolution See Punctuationism
Species (biological) A reproductively isolated aggregate of populations
which can interbreed with one another becau se they share the san1e isolat­
ing mechanis m s . See Evolu tionary species concept; Nominalist species
concept; Typological species con cept
Species replacement The tu rnover of species in tin1 e, owing to the ex­
tinction of some species and the o rigin of new species that replace them .
Also referred to as species selection
Sport In animal b reedin g , an aberrant individual produced by a maJor
m u tation
Stochastic processes Processes consisting of a senes of steps , each of
which is rando m in directio n . See Determinism
S y mpatric speciation The splitting of a species into two reproductively
isolated species within the same area
S y mpatric Coexisting at the same locality ; said of a population in breed­
ing condition that exists within the cruising range of individuals that be­
long to a different species
S y stematics The science which studies the diversity of organis ms
Taxon A monoph yletic group of organisn1s that share a definite set o f
cha racters and are considered sufficiently distinct to be wo rthy o f a forn1a l
name
Taxonom y The theory and practice o f classifying organisn1 s
Teleolog y The actual or only seen1ing exis tence o f end-d irected pro­
cesses in nature, and their study. See Finalis m

• 1 86 .
 Glossa ry

Teleomatic p rocess A seemingly end-directed process that is strictly

con trolled by natural laws such as the law of gravity or the firs t law of
thermodyn a n1ics
Teleonomic p rocess A process or behavior that owes its goal­
directedness to the o pera tion o f a program
Theism The belief in a personal god who is forever present and able to
affect any natu ral process a t any time
Transformational evolution G radual change of an obj ect in the course
of time from one condition of existence into another; usually co mbined
with a belief in change fro m " l ower " to " higher, " or from less perfect to
more perfect . See L a ma rckis m ; Saltational evolution ; Va riational evol u tion
Ty p ological s p ecies conce p t Defines species on the basis of degree o f
difference . See Biological species concept; Evolutionary species con cep t;
N o m inalist species concept
Ultimate or evolutionar y causation In evolutiona ry biology, the his­
to rical factors responsible for the properties of species and individuals , and
more specifically for the com position of the genotype
Uniformitarianism A theory, particularly p romoted by Charles Lyell ,
tha t all geological changes , no m a t ter at w h a t rate they occur, are gradual .
See C a tastrophism
Variant A m ember o f a variable population
Varia-t ional evolution Darwin's concept of evolutio n , which states that
change occurs in every generation through the p roduction of a large
a m ount of new genetic variation and through the survival (" selection " ) o f
a s mall percentage of the varied individuals t o serve a s the progenitors o f a
next generatton . See Sal tational evolu tion ; Selection ; Transformational
evolution
Yarrell 's Law A generalization proposed by William Yarrell that " the
longer a character was in the blood of a race, the more fi xed it would be"
Z y gote A fertilized egg; the indi vidual that results fro m the union of two
ga metes and their n u clei
.. ,
 A C K N O W L E D G M E N T S

This anal ysis and evalua tion of Darwin's though t is based not only
on Darwin's own writings but also on the critical studies of a ded­
ica ted group of Da rwin scholars . To all of then1 I a n1 greatly in­
debted . Their n u m ber is too large for me to men tion them here
in dividually, b u t their na mes can be found in the Referen ces . A t
t h e r i s k of bein g u n fair to t h e o thers , I wan t t o thank es pecially
Frederick B u rk h a rd t and David Kohn fo r their never-failing will­
ingness to g i ve m e s u ggestions and information .
Walter B o ra wski again indefatigably typed cou n tless drafts of
the manu scri p t and assisted me in the preparation of the Refer­
en ces , Glossa ry, and Index . Howard Boyer, Senior Science Editor
at Ha rva rd Univers ity Pres s , encou raged and supported this pro-
ject fr o m its inception . Susan Wal lace , also at the Pres s , i m proved
the ra w m an uscript i m measurably by eliminating redundan cy, ar­
ran gi n g a m o re logical sequence of subject matter, pointing out
gaps in m y original manus cri p t , sugges ting stylistic i m p rove­
men t s , and in various o ther ways. I cannot express in words my
deep gratitu de to her.

Ernst Mayr
..
 Index

Adams, M . , 1 3 9 Bowler, P. J . , 5 5 , 60
adaptatio n , 5 6, 1 1 5 Boyle, R . , 5 2
adaptationist progra m , 1 1 5 branching, 2 1 , 2 2
adap tive radiations, 1 5 7 " bridgeless gaps , " 4 1
a ffi n ity, 2 2 Buch , L . v . . 1 9 , 3 2
Agassiz, L . , 8 , 1 6 , 3 8 , 1 oo Buffon, G . L . , 1 2 , 5 1
A lexander, R . , 1 5 5 Burian, R . , 9 1
altruis m , 1 5 5
a m ph i m i xis , 1 2 4 Candolle, A . P. de, 78
animal breeders , 7 1 , 8 1 causation s : p ro x i mate, 5 3 ; u l t i mate, 5 3 ;
an ti-crea tion is m , 1 4 1 s i m u l taneou s , 1 4 7
A rchaeop teryx , 1 5 9 causes , fi na l , 50
a rchetypes, 5 5 cent ral dog m a , 1 20
u argu men t , one lon g , " 94 chan ce, 49, 66, 1 40 ; or necessity, 59
A ri s totle, 50 change. 1 5 3
asexuality, adva n tage o f, 1 23 cha ractcr divergence, 3 3 , 3 6
autobiography ( Darwin's) , 70 Chardin, T. de, 66
Chetverikov, S . , 1 3 4
Baconian method, 9 chorda . 24
Baer, K . E . v . , 5 3 Christian faith, 7 5
balance o f nature , 7 5 Churchill, F. , 1 1 9, 1 2 1
balanced selectio n , 1 5 1 classi fica tion , theory o f, 2
barnacles , 6 , 45 cohesion , of genes , 1 60
Barzu n , J . , 1 0 8 com mon descen t , 2 1 , 2 2 , 9 5 , 1 00 , 1 6 3
Bates, H . W. , 8 com petitio n , 5 8 , 78
Bateson , W. , 46 consnousness, 2 5
Baup lan , 1 60 cons train ts, 1 29 ; ideologica l , 3 8 ; devel-
Bau r, E . , 1 3 3 opmen t a l , 1 I 6, 1 64
bdelloid rotifers, I 2 3 con tinental drift , 1 o 5
Bea_R/e, 4 , 69 crea t10msm , 1 5 , 1 6, 1 7 , 99
Beckn er, M . , 66 C reator, 49
beh avior, social , 1 54 cu lt u re, 63
Bergson , H . , 66 C u vier, G . , 1 6 , 2 2 , 77
Bible, 1 3
Bla cher, L. I . , I 1 9 Darwin , A nnie, I 5
Blu men bach , J . F. , r 6 Darwin , Cha rles : as agnostic, I 3 - 1 5 ;
Blyth, E . , 86 fi ve theories of, J 5 ; a s phi losopher.
Bon net, C . , 78 5 0 , 1 0 2 ; papers o f, 7 I

• 191 •
 Index
.'
Darwin, E m m a , 1 5 exemplar, 1 06
Darwin, E rasmus, 3 experi ment, Darw in's use of. 1 0
Darwinian revo lution, 89, 1 07, 1 3 2 ; explanatory model, Darwin's . 7 3
fi rst, 1 2 , 2 5 ; second, 89 extern alis m , 39
Darwinis m , 90; as anticreationis m , 93 ; extinction , I 6, 5 5
as evolutionism , 9 3 ; a s anti­ eye, evolu tion of. 5 8
ideology, 96; as selectionism, 97; as
variational evolution, 97; as creed of fe male choice. 8 9 , 1 1 7
the Darwinians, 98; as a new wo rld­ finalism, 50-67
view, 1 o 1 ; as a new methodology, Fisher, R. A . , 1 43 , 1 5 1
1 04 ; current meaning of. 1 07, 1 62 , Fitz Roy, R . , 4
1 64 ; stages of, 1 4 3 Ford. E . B . , 1 3 4
de Beer, G . , 7 1 fossil record, 1 6, 5 5 . I 3 8
de V ries, H . , 46 founder popula tions , 1 4 8
dei s m , 93
Derh a m , W. , 5 2 Galapagos, 5
Descartes, R . , 5 1 Gal con , F. , 4 5 , 1 1 9
Descent of Man , 7 game theory, 1 56
design, 3 8 , 5 2 , 5 4 gene flow, 1 4 8
determinants, genetic, 1 2 5 genetic code, 2 3 , 1 6 3
determinism, 48 genetic drift. 1 1 9
developmen t , 1 4 5 , 1 5 8 , 1 62 genetic prog ra m, 1 2 2
diagra m, branching, 22 genetic variation, sou rce of. 1 24
dinosaurs , 6 5 genet1nsrs. 1 3 9
diversification . 62 genetics, 1 3 2 , I 3 9
diversity, 1 6 3 genotype: do mains of. 1 5 8 : cohesion
Dobzhansky, Th . , 1 3 4 of, 1 3 0, 1 60; con gealing of. 1 60
domestication , 44 Geotfroyis m . 1 26
dominance, 63 geographical speciation, 20
geology, 1 6
earth. age of, 1 6 germ plas m , 1 22
East, E . , 1 3 3 germinal selection , 1 2 5 , 1 26
Ehrenberg, C . G . . 76 Ghiselin, M . T. , 6, 4 2 , 79 , 1 0 5
E i mer. T. , 6 1 gill arches . 24
Eldredge. N . , 1 3 7 Gillespie, N . C . , 1 5
electrophoresis, 1 5 2 Gillis pie. C . , 94
environment, emancipation fro m , 63 God. 5 1 . 5 2 , 56, 96
equilibri a . punctuated, I 5 3 Goldsch midt. R . . 46
essen tialis m , 40 Good man, M . . 1 59
euka ryotes . 62 Gordon. S . . 8 5 . 1 02
evolu tion : g radual , 1 8 , 43 ; saltational. Gould. J . , 5 . 1 8 , 3 2
42 ; transform a tion a l , 43 : variational . Gould . S . J . , 1 1 2. 1 3 7
4 3 . 1 6 3 : unpredictable. 44; mecha­ gradualis m , 1 7 , 1 8 , 4 4 . 1 1 4. 1 6 3
nism o f. 68 : as fact, 1 1 2. 1 62 ; geo­ Gran t , V. . 1 4 5
graphica l , 1 3 7; vertica l . 1 3 7 Gray. A . , 5 9. 1 oo
evolu tionary synthesis. 1 06, 1 3 2- 1 40. Greene. J . C . , 96 . 1 o 1
1 4 1 , q6 group. cul tural, 1 5 6
evolu tion ism : vertica l . I 7, 20; horizon­ group selection. 1 5 6 . 1 5 7
ta l . 20 Gruber. H . E . . 4 3 . 7 1 . 80

• 1 92 .
 Index

Haeckel , E . . 2 4 , 60 , 1 5 7 K lcinen berg , N . , 1 6 1

Haldare , J . B . S . . 1 5 5 Kolliker, A . , 46, 59
H a m i l ton, W. D . . 1 5 4 Kohn , D . . 1 4 , 1 5 , 44, 6 9, 70, 73 , 1 08
Harris . H . , 1 5 2 K u hn, T. , 1 3 2
Hartman n . E . v . . 5 3
Henslow. J . S . . 5 5 Lamarck , ] . B . , 1 2 , 1 6 . 1 7 , 1 9, 2 1 , 4 3 ,
Herbert, S . , 79 54, 62
Herder, J . G . . 5 2 Lamarck is m , 1 20
Herschel . J . , 9, 4 8 , 5 6 laws, 49 , 5 6 , 5 7 , 9 3
Hessen , B . , 3 9 Leibni z . G . W. . 4 5 , 5 2
higgledy-pig gledy, theory o f the, 49 Lenno x . J. G . . 5 3
H i m melfarb, G . , 1 08 Lenoir, T. , 50
Hodge , J. , 69, 9 5 Lewontin , R. C . . 43 , 1 5 2
Holbach. B a ron , 5 1 L i moges , C . , 7 1
holis m , 1 3 8 Linnacan hierarch y, 2 3
ho meoboxes, 1 5 8 Linnaeus, C . , 20, 7 8
ho moplasy. 1 5 I Low, R . , 5 3
Hooker, J . , 7 Lovejoy, A . , 44
horizontal evo l u tion. 20, 1 3 7 lun g fi shes , 1 6 I
Hubby, J . L . , 1 5 2 L wotf, A . , 1 3 9
Hu 11. D . L . , I 4 ' 9 I , I 06 Lyell , C . , 4 , 1 2 , 1 6, 1 7 , 1 9 , 3 8 , 4 1 , 43 ,
Hume, D . , 5 4 5 5 , 64, I O I
Hu xley, A . , 4 7 Lysenko, 1 3 9
Huxley, J . . 1 3 4
H u xley, T. H . , 8 , 24, 46, 6 3 ma croevolution , I 3 8 , 1 6 1
Malthus, R . , 5 7, 69, 8 1 , 84- 8 5 , 1 02
ideologies , 3 8 ; opposin g Da rwinis m , m a n : posi tion of, 24, 3 8 -3 9; origin of,
40 25
inclusive fi tness, 1 5 6 ma tter in motion, 96
individual , 8 0 ; uniq ueness of, 4 2 Maynard S mith, J. , 1 5 6
inductionis m , 1 2 7 mechanists, 5 0
inductivis m , 1 04 Men de l , G . , 1 09
inferen ce, 4 8 , 1 08 Mendelians, 46, I 3 6
inheritance : soft , 74, 1 08 ; of acquired Mendel 's la ws, 1 49
characters, 74, 1 1 o, 1 1 8 ; blendin g , method , scien tific, 9
1 09 , 1 2 3 ; particulate, 1 29, 1 3 5 methodology, 1 04- 1 0 5
intern a l causation , 69 M i l l , J . S . , 9, 4 1 , 56
mind, 2 5
Jacob, F. , 1 49 mockingbirds, 5 , 1 8 , 20, 2 2 , 3 2
Jen kin, F. , 46 molecu lar biology, 1 49
Jou rnal of Research es , 4 molecu lar clock , 1 5 9
Jukes , T. , 1 52 M o n o d , J . , 66
Moore, J. R . , I 5 , 59
K an t , I . , 5 0 , 54 Morgan, T. H . , 60, I 3 3
Kellogg, V. L . . 60 mosaic evolution, 1 3 0, 1 5 5 , 1 5 8
K i mura , M . , 1 5 2 M u l ler, F. , 8
k i n selection , 1 56 M u l ler, H . J. . I 5 I
King, J . , 1 52 mu tation , spontaneou s, I 24
K i tcher, P. , 9 1 , 9 8 , 1 0 5 muta tion s , s m a ll, 1 3 3

• 1 93 .
 Index
.'
Nagel , E. , 66 progress, 1 , 5 3 , 5 7; evolutionary, 62. 64
natural kinds, 27, 4 1 progressionists, 4 3
natural selection, 5 7 , 59, 6 5 , 70, 86, progressi veness , 6 3
I I 3 , I 3 9, I 64; path to, 68; cause of, punctuationi s m , I 5 3
86; validity of, 1 2 1 ; triu m ph of, 1 3 9.
See also Selection quality, 1 2 5
natural theology, 3 8 , 5 2 , 5 5 , 68 qumanans, 22
naturalist, Darwin as, 3 , 1 o, 1 3 9
naturalists, con tribu tions by, 1 3 7- 1 3 9 rate of evolu tion , 4 7, I 59
neo-Darwini s m , 1 08 Ray, J. , 20, 52
neo-Lamarck i s m , 60 recapitulation, 1 1 2 , 1 4 5 , I 57
nervous system , central, 63 Recker, D . , 95, 99, 1 0 1 , 1 04
neutral evolution , 1 5 1 recombination , genetic, 1 2 2, 1 24
notebooks (Darwin), 8 3 -84 rectilinearity, 6 1 , 6 5
novelties , evolu tionary, 1 3 8 reduction division , 1 29
rel igion, Da rwin's, 1 3
organs , vestigial, 5 7 Rensch , B . , I 3 4
origin of life, 2 4 reproduction, sexual, 8 2
orthogenesis, 5 3 , 60, 1 13 resources, limitation 'o f, 76
Osborn, H . F. , 43 revolution : first Da rwin ian . 1 2 , 2 5 ; sec-
Ospova t, D . , 1 5 , 56 ond Darwinian , 89
Owen , R . , 22 revolution s : in tel lectua l . 1 ; scien tific, 3 9
O xford, I 3 9 Rhea , 1 8
Richards , R . J. , 1 5
pan genesis , 3 6 Ro manes , G . J . , 1 1 0
Patagonia, 4 Ruse, M . . I 3 , 56, 8 2
pathogens, 8 7 Russia, I 3 9
perfecting principle, 6 1
perfection . 5 2 , 5 5 , 5 8 ; progression to- Sachs, J. , 6 1
wa rd , 56 sal tation, 1 8 , 46
phenotype, I 3 7 sal tationis m , 1 9, 42 . I 1 3
philosophers , 4 8 , 50 Sap p , J. , I 3 6
philosophy : of science, 2 , 9; hy pothe- sea /a natu rae, 2 1 . 6 I
tico-deductive, 1 2 8 Schindewolf, 0 . , 46
phylogeny, 1 1 2 Schleiden, M . . 5 5
physicalism . 48 Sedg wick , A . . 5 5
physicists, 48 selection : artifi cial . 8 3 ; of 8 7 ; for, 8 8 ;
Plato , 40 sexual. 8 8 , 1 1 7 , 1 6 1 ; evidence for.
plurali s m , 1 0 5 , 1 06, 1 47 1 1 4; normalizin g , 1 1 6; target of,
population stability, 76 1 1 7 ; germinal, 1 2 5 . Sa also Na tural
population thinking, 40, 4 2 , 46, 74, 80, selection
138 se x . signi ficance of. 1 22. 1 24
popula tion s : local. 2 7; ex ponential in- sexual selection. 8 8 . 1 1 7. 1 ,6. 1 64
crease o f. 7 5 sibling species . 27. 1 4 8 . 1 6 1
prediction . 48 Sigwart, C . . 66
principle of exclusion , 1 1 6 Sim pson . G . G . . 1 8 . 6 5 , 1 3 5 . 1 5 9
probabilis m , 49 Smith. A . . 1 02
processes : stochastic. 49; teleomatic. 67 S mith. S . . 7 1
progra m : genetic, 1 5 8 ; somatic. 1 6 1 sociobiology. 1 5 4- 1 5 7

• 1 94 .
 Index

socioecono m ic factors. in scientific rev- twin studies, 1 5 4

ol utions . 3 9 typologica l thinking. 4 2
somatic progra m s , 1 5 8
sou l . in m a n . 3 9 uniformitaria nis m . 1 6, 4 3
special creation . 1 02 uniqueness. 74
specia tion. 5 . 1 8 , 1 9; a l lopatric, 20; de­ use and disuse, ] 6. 1 09 , I 20
fined , 3 1 - 3 4 ; g radu a l , 3 2 ; on isl ands.
van Bcneden , E . . I 2 3
3 2 ; s y m pa t ric. 3 2
va riation . 4 2 , 1 09; accidental. 49; sou rec
species concepts , 26; nom inalist. 2 7 ;
of, 1 1 8 ; q anti tat i vc . 1 2 5 ; g rad u a 1 .
typologica l . 2 7 ; evolutiona ry. 2 8 ;
1 36
biologica l . 29; Darwin's . 3 0
variety. 29
species replacement , 8 8
ve rtical evolution . 1 7 . 20. 1 3 7
.. species book , '' 6
Voltaire, F. M . A rouet. 5 4
species : co nstancy of. 1 2 ; defined . 27 �
30 ; in plants. 29; m u l tiplication o t . Wa ddington. C . H . . I 5 7
3 1 ; inci pien t , 3 3 Wa gner. M . , 1 9 . 3 2
spec ies selection . 64 . 8 8 Wallace. A . R . , 6 , 1 9 , 7 3 , 90. 92. 97
species su ccession. 64 Wedgwood , E . , 6
Spen cer. H . , 1 02 . 1 2 5 Wedgwood. J. . 3
Stanl ey. S . M . . 4 3 Wegener, A . . 1 2
sta sis. 64 , 6 5 Weisman n , A . , 60 , 1 1 o. 1 o 8 - 1 3 1 ; as
Stebbins, G . L . . I 3 4 , 1 4 5 pansclection ist. 1 1 6 ; criticism of.
stochastic processes . 1 40 1 2 7 ; con tributions of. 1 2 8
struggle for e xistence. 77 Westol l . S . . 1 6 1
S u l loway. F. J. . 5 . 1 8 . 3 4 Whewel l. W. . 9, 4 1 , 56
syn thesis. evolutiona ry. I 06 , 1 3 2- 1 40. White, M. J. D . . 1 4 5
1 .p . 1 4 6 Whitehead. A . N . . 66
system a tics . 1 46 Whitman . C . 0 . . 6 1
systems. 97; ada pted , 67 .. why " question s, 4
Wi ley. E . 0 . . 2 8
target of select ion, 68 Willis. J. C . . 46
teleology. 50, 66 ; cos mic, 5 3 . 67 Wi l l m ann. R . , 2 8
tclco m atic processes . 67 Wi lson . A . C . . 1 5 9
teleonomic processes. 67 Wi lson . E . 0 . . 1 54
theis m , 1 2
To u l m i n , S . . 5 3 Ya rrcl l 's L a w. 44
transmutation , 4 3
Trivcrs , R . . 1 5 5 Zfitgl'ist , 1. 40

• 1 95 •
 .. ,

L ibra ry of Co11.�ress Ca ta lo,� i11,�- i11 - Publica tio11 Data

Mayr. Ernst . 1 904-

0ne long argu ment : Charles Darwin and the genes is of modern
evolu tionary thought / Ernst Mayr.
p. cm .
Includes bibliographical re ferences and inde x .
I S B N 0-6 74-6 3 905-7 (cloth)
I S B N 0-674-6 3906- 5 (paper)
I . Evol u tion . 2 . Evolution- Phi losophy.
3 . Darwin. Charles . 1 809 - 1 8 8 2 . I . Ti tk.
QH J 7 1 . M J 3 6 1 99 1
, 7 , -dc20
91-1 105 I
CIP