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Also by Richard Dawkins

The Selfish Gene

The Extended Phenotype
The Blind Watchmaker
River Out of Eden
Climin! "ount #mproale
$n%eavin! the Raino%
& 'evil(s Chaplain
The &ncestor(s Tale
The God 'elusion
T)E E+#'E*CE ,OR E+O-$T#O*
.osh Timonen
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Chapter 2 Only a theoryB
Chapter 3 'o!s0 co%s and caa!es
Chapter 4 The primrose path to macro:evolution
Chapter > Silence and slo% time
Chapter = Before our very eyes
Chapter < "issin! linkB What do you mean0 (missin!(B
Chapter @ "issin! personsB "issin! no lon!er
Chapter ; 6ou did it yourself in nine months
Chapter 9 The ark of the continents
Chapter 25 The tree of cousinship
Chapter 22 )istory %ritten all over us
Chapter 23 &rms races and (evolutionary theodicy(
Chapter 24 There is !randeur in this vie% of life
&PPE*'#C The history:deniers
B#B-#OGR&P)6 &*' ,$RT)ER RE&'#*G
T)E evidence for evolution !ro%s y the day0 and has never een stron!er1 &t the same
time0 paradoxically0 ill:informed opposition is also stron!er than # can rememer1 This ook is my
personal summary of the evidence that the (theory( of evolution is actually a fact : as
incontrovertile a fact as any in science1
#t is not the first ook # have %ritten aout evolution0 and # need to explain %hat(s different
aout it1 #t could e descried as my missin! link1 The Selfish Gene and The Extended Phenotype
offered an unfamiliar vision of the familiar theory of natural selection0 ut they didn(t discuss the
evidence for evolution itself1 "y next three ooks0 in their different %ays0 sou!ht to identify0 and
dissolve0 the main arriers to understandin!1 These ooks0 The Blind Watchmaker0 Rier !"t of
Eden and 7my favourite of the three8 #limbin$ %o"nt &mprobable0 ans%ered Euestions like0 (What
is the use of half an eyeB( (What is the use of half a %in!B( ()o% can natural selection %ork0 !iven
that most mutations have ne!ative effectsB( Once a!ain0 ho%ever0 these three ooks0 althou!h they
cleared a%ay stumlin! locks0 did not present the actual evidence that evolution is a fact1 "y
lar!est ook0 The Ancestor's Tale0 laid out the full course of the history of life0 as a sort of ancestor:
seekin! Chaucerian pil!rima!e !oin! ack%ards in time0 ut it a!ain assumed that evolution is true1
-ookin! ack on those ooks0 # realiFed that the evidence for evolution itself %as no%here
explicitly set out0 and that this %as a serious !ap that # needed to close1 The year 3559 seemed like a
!ood time0 it ein! the icentennial year of 'ar%in(s irth and the 2=5th anniversary of !n the
!ri$in of Species1 *ot surprisin!ly0 the same thou!ht occurred to others0 and the year has seen some
excellent volumes0 most notaly .erry Coyne(s Why Eol"tion is Tr"e1 "y hi!hly favourale revie%
of his ook in the Times (iterary S"pplement is reproduced at
The %orkin! title under %hich my literary a!ent0 the visionary and indefati!ale .ohn
Brockman0 offered my ook to pulishers %as !nly a Theory1 #t later turned out that Denneth
"iller had already pre:empted that title for his ook:len!th response to one of those remarkale
courtroom trials y %hich scientific syllauses are occasionally decided 7a trial in %hich he played a
heroic part81 #n any case0 # had al%ays douted the title(s suitaility for my ook0 and # %as ready to
shelve it %hen # found that the perfect title had een lurkin! on another shelf all alon!1 Some years
a!o0 an anonymous %ell:%isher had sent me a T:shirt earin! the BarnumesEue slo!anA (Evolution0
the Greatest Sho% on Earth0 the Only Game in To%n(1 ,rom time to time # have %orn it to !ive a
lecture %ith that title0 and # suddenly realiFed that it %as ideal for this ook even if0 in its entirety0 it
%as too lon!1 # shortened it to The Greatest Show on Earth1 (Only a Theory(0 %ith a precautionary
Euestion mark to !uard a!ainst creationist Euote:minin!0 %ould do nicely as the headin! to Chapter
# have een helped in various %ays y many people0 includin! "ichael 6udkin0 Richard
-enski0 Geor!e Oster0 Caroline Pond0 )enri '1 Grissino:"ayer0 .onathan )od!kin0 "att Ridley0
Peter )olland0 Walter .oyce0 6an Won!0 Will &tkinson0 -atha "enon0 Christopher Graham0 Paula
Diry0 -isa Bauer0 O%en Selly0 +ictor ,lynn0 Daren O%ens0 .ohn Endler0 #ain 'ou!las:)amilton0
Sheila -ee0 Phil -ord0 Christine 'eBlase and Rand Russell1 Sally Gaminara and )ilary Redmon0
and their teams in 7respectively8 Britain and &merica0 have een %onderfully supportive and can:
do:ish1 On three occasions %hile the ook %as !oin! throu!h the final sta!es of production0 excitin!
ne% discoveries %ere reported in the scientific literature1 Each time0 # diffidently asked if the
orderly and complex procedures of pulication mi!ht e violated to accommodate the ne% find1 On
all three occasions0 far from !rumlin! at such disruptive last:minutemanship0 as any normal
pulisher mi!ht0 Sally and )ilary !reeted the su!!estion %ith cheerful enthusiasm and moved
mountains to make it happen1 EEually ea!er and helpful %as Gillian Somerscales0 %ho copy:edited
and collated the ook %ith literate intelli!ence and sensitivity1
"y %ife -alla Ward has once a!ain sustained me %ith unfailin! encoura!ement0 helpful
stylistic criticisms and characteristically stylish su!!estions1 The ook %as conceived and e!un
durin! my last months in the professorship that ears the name of Charles Simonyi0 and completed
after # retired1 #n si!nin! off as Simonyi Professor0 fourteen years and seven ooks after our
momentous first meetin!0 # %ould once a!ain like to express my !rateful appreciation to Charles1
-alla Hoins me in hopin! that our friendship %ill lon! continue1
This ook is dedicated to .osh Timonen0 %ith thanks to him and to the small and dedicated
and %ho ori!inally %orked %ith him to set up Richard'a%kins1net1 The %e kno%s .osh as an
inspired site desi!ner0 ut that is Hust the tip of an amaFin! iceer!1 .osh(s creative talent runs deep0
ut the ima!e of the iceer! captures neither the versatile readth of his contriutions to our Hoint
endeavour0 nor the %arm !ood humour %ith %hich he makes them1
O*-6 & T)EOR6B
# "&G#*E that you are a teacher of Roman history and the -atin lan!ua!e0 anxious to
impart your enthusiasm for the ancient %orld : for the ele!iacs of Ovid and the odes of )orace0 the
sine%y economy of -atin !rammar as exhiited in the oratory of Cicero0 the strate!ic niceties of the
Punic Wars0 the !eneralship of .ulius Caesar and the voluptuous excesses of the later emperors1
That(s a i! undertakin! and it takes time0 concentration0 dedication1 6et you find your precious
time continually preyed upon0 and your class(s attention distracted0 y a ayin! pack of i!noramuses
7as a -atin scholar you %ould kno% etter than to say (i!norami(8 %ho0 %ith stron! political and
especially financial support0 scurry aout tirelessly attemptin! to persuade your unfortunate pupils
that the Romans never existed1 There never %as a Roman Empire1 The entire %orld came into
existence only Hust eyond livin! memory1 Spanish0 #talian0 ,rench0 Portu!uese0 Catalan0 Occitan0
RomanshA all these lan!ua!es and their constituent dialects spran! spontaneously and separately
into ein!0 and o%e nothin! to any predecessor such as -atin1 #nstead of devotin! your full attention
to the nole vocation of classical scholar and teacher0 you are forced to divert your time and ener!y
to a rear!uard defence of the proposition that the Romans existed at allA a defence a!ainst an
exhiition of i!norant preHudice that %ould make you %eep if you %eren(t too usy fi!htin! it1
#f my fantasy of the -atin teacher seems too %ay%ard0 here(s a more realistic example1
#ma!ine you are a teacher of more recent history0 and your lessons on t%entieth:century Europe are
oycotted0 heckled or other%ise disrupted y %ell:or!aniFed0 %ell:financed and politically muscular
!roups of )olocaust:deniers1 $nlike my hypothetical Rome:deniers0 )olocaust:deniers really exist1
They are vocal0 superficially plausile0 and adept at seemin! learned1 They are supported y the
president of at least one currently po%erful state0 and they include at least one ishop of the Roman
Catholic Church1 #ma!ine that0 as a teacher of European history0 you are continually faced %ith
elli!erent demands to (teach the controversy(0 and to !ive (eEual time( to the (alternative theory( that
the )olocaust never happened ut %as invented y a unch of Iionist faricators1 ,ashionaly
relativist intellectuals chime in to insist that there is no asolute truthA %hether the )olocaust
happened is a matter of personal eliefJ all points of vie% are eEually valid and should e eEually
The pli!ht of many science teachers today is not less dire1 When they attempt to expound
the central and !uidin! principle of iolo!yJ %hen they honestly place the livin! %orld in its
historical context : %hich means evolutionJ %hen they explore and explain the very nature of life
itself0 they are harried and stymied0 hassled and ullied0 even threatened %ith loss of their Hos1 &t
the very least their time is %asted at every turn1 They are likely to receive menacin! letters from
parents0 and have to endure the sarcastic smirks and close:folded arms of rain%ashed children1
They are supplied %ith state:approved textooks that have had the %ord (evolution( systematically
expun!ed0 or o%dleriFed into (chan!e over time(1 Once0 %e %ere tempted to lau!h this kind of
thin! off as a peculiarly &merican phenomenon1 Teachers in Britain and Europe no% face the same
prolems0 partly ecause of &merican influence0 ut more si!nificantly ecause of the !ro%in!
#slamic presence in the classroom : aetted y the official commitment to (multiculturalism( and the
terror of ein! thou!ht racist1
#t is freEuently0 and ri!htly0 said that senior cler!y and theolo!ians have no prolem %ith
evolution and0 in many cases0 actively support scientists in this respect1 This is often true0 as # kno%
from the a!reeale experience of collaoratin! %ith the then Bishop of Oxford0 no% -ord )arries0
on t%o separate occasions1 #n 355> %e %rote a Hoint article in the S"nday Times %hose concludin!
%ords %ereA (*o%adays there is nothin! to deate1 Evolution is a fact and0 from a Christian
perspective0 one of the !reatest of God(s %orks1( The last sentence %as %ritten y Richard )arries0
ut %e a!reed aout all the rest of our article1 T%o years previously0 Bishop )arries and # had
or!aniFed a Hoint letter to the then Prime "inister0 Tony Blair0 %hich read as follo%sA
'ear Prime "inister0
We %rite as a !roup of scientists and Bishops to express our concern aout the teachin! of
science in the Emmanuel City Technolo!y Colle!e in Gateshead1
Evolution is a scientific theory of !reat explanatory po%er0 ale to account for a %ide ran!e
of phenomena in a numer of disciplines1 #t can e refined0 confirmed and even radically altered y
attention to evidence1 #t is not0 as spokesmen for the colle!e maintain0 a (faith position( in the same
cate!ory as the ilical account of creation %hich has a different function and purpose1
The issue !oes %ider than %hat is currently ein! tau!ht in one colle!e1 There is a !ro%in!
anxiety aout %hat %ill e tau!ht and ho% it %ill e tau!ht in the ne% !eneration of proposed faith
schools1 We elieve that the curricula in such schools0 as %ell as that of Emmanuel City
Technolo!y Colle!e0 need to e strictly monitored in order that the respective disciplines of science
and reli!ious studies are properly respected1
6ours sincerely
The Rt Red Richard )arries* Bishop of !xford+ Sir Daid Attenboro"$h ,RS+ The Rt Red
#hristopher )erbert* Bishop of St Albans+ (ord %ay of !xford* President of the Royal
Society+Professor -ohn Enderby ,RS* Physical Secretary* Royal Society+ The Rt Red -ohn !lier*
Bishop of )ereford+ The Rt Red %ark Santer* Bishop of Birmin$ham+ Sir .eil #halmers* Director*
.at"ral )istory %"se"m+ The Rt Red Thomas B"tler* Bishop of So"thwark+ Sir %artin Rees ,RS*
Astronomer Royal+ The Rt Red /enneth Steenson* Bishop of Portsmo"th+ Professor Patrick
Bateson ,RS* Biolo$ical Secretary* Royal Society+ The Rt Red #rispian )ollis* Roman #atholic
Bishop of Portsmo"th+ Sir Richard So"thwood ,RS+ Sir ,rancis Graham0Smith ,RS* Past Physical
Secretary* Royal Society+ Professor Richard Dawkins ,RS
Bishop )arries and # or!aniFed this letter in a hurry1 &s far as # rememer0 the si!natories to
the letter constituted 255 per cent of those %e approached1 There %as no disa!reement either from
scientists or from ishops1
The &rchishop of Canterury has no prolem %ith evolution0 nor does the Pope 7!ive or
take the odd %ole over the precise palaeontolo!ical Huncture %hen the human soul %as inHected80
nor do educated priests and professors of theolo!y1 This is a ook aout the positive evidence that
evolution is a fact1 #t is not intended as an anti:reli!ious ook1 #(ve done that0 it(s another T:shirt0
this is not the place to %ear it a!ain1 Bishops and theolo!ians %ho have attended to the evidence for
evolution have !iven up the stru!!le a!ainst it1 Some may do so reluctantly0 some0 like Richard
)arries0 enthusiastically0 ut all except the %oefully uninformed are forced to accept the fact of
evolution1 They may think God had a hand in startin! the process off0 and perhaps didn(t stay his
hand in !uidin! its future pro!ress1 They proaly think God cranked the universe up in the first
place0 and solemniFed its irth %ith a harmonious set of la%s and physical constants calculated to
fulfil some inscrutale purpose in %hich %e %ere eventually to play a role1 But0 !rud!in!ly in some
cases0 happily in others0 thou!htful and rational churchmen and %omen accept the evidence for
What %e must not do is complacently assume that0 ecause ishops and educated cler!y
accept evolution0 so do their con!re!ations1 &las0 as # have documented in the &ppendix0 there is
ample evidence to the contrary from opinion polls1 "ore than >5 per cent of &mericans deny that
humans evolved from other animals0 and think that %e : and y implication all of life : %ere created
y God %ithin the last 250555 years1 The fi!ure is not Euite so hi!h in Britain0 ut it is still
%orryin!ly lar!e1 &nd it should e as %orryin! to the churches as it is to scientists1 This ook is
necessary1 # shall e usin! the name (history:deniers( for those people %ho deny evolutionA %ho
elieve the %orld(s a!e is measured in thousands of years rather than thousands of millions of years0
and %ho elieve humans %alked %ith dinosaurs1 To repeat0 they constitute more than >5 per cent of
the &merican population1 The eEuivalent fi!ure is hi!her in some countries0 lo%er in others0 ut >5
per cent is a !ood avera!e and # shall from time to time refer to the history:deniers as the (>5:
To return to the enli!htened ishops and theolo!ians0 it %ould e nice if they(d put a it
more effort into comatin! the anti:scientific nonsense that they deplore1 &ll too many preachers0
%hile a!reein! that evolution is true and &dam and Eve never existed0 %ill then lithely !o into the
pulpit and make some moral or theolo!ical point aout &dam and Eve in their sermons %ithout
once mentionin! that0 of course0 &dam and Eve never actually existedK #f challen!ed0 they %ill
protest that they intended a purely (symolic( meanin!0 perhaps somethin! to do %ith (ori!inal sin(0
or the virtues of innocence1 They may add %itherin!ly that0 oviously0 noody %ould e so foolish
as to take their %ords literally1 But do their con!re!ations kno% thatB )o% is the person in the pe%0
or on the prayer:mat0 supposed to kno% %hich its of scripture to take literally0 %hich
symolicallyB #s it really so easy for an uneducated church!oer to !uessB #n all too many cases the
ans%er is clearly no0 and anyody could e for!iven for feelin! confused1 #f you don(t elieve me0
look at the &ppendix1
"I still say it's only a theory."
Think aout it0 Bishop1 Be careful0 +icar1 6ou are playin! %ith dynamite0 foolin! around
%ith a misunderstandin! that(s %aitin! to happen : one mi!ht even say almost ound to happen if
not forestalled1 Shouldn(t you take !reater care0 %hen speakin! in pulic0 to let your yea e yea and
your nay e nayB -est ye fall into condemnation0 shouldn(t you e !oin! out of your %ay to counter
that already extremely %idespread popular misunderstandin! and lend active and enthusiastic
support to scientists and science teachersB
The history:deniers themselves are amon! those that # am tryin! to reach in this ook1 But0
perhaps more importantly0 # aspire to arm those %ho are not history:deniers ut kno% some :
perhaps memers of their o%n family or church : and find themselves inadeEuately prepared to
ar!ue the case1
Evolution is a fact1 Beyond reasonale dout0 eyond serious dout0 eyond sane0 informed0
intelli!ent dout0 eyond dout evolution is a fact1 The evidence for evolution is at least as stron! as
the evidence for the )olocaust0 even allo%in! for eye %itnesses to the )olocaust1 #t is the plain
truth that %e are cousins of chimpanFees0 some%hat more distant cousins of monkeys0 more distant
cousins still of aardvarks and manatees0 yet more distant cousins of ananas and turnips 1 1 1
continue the list as lon! as desired1 That didn(t have to e true1 #t is not self:evidently0
tautolo!ically0 oviously true0 and there %as a time %hen most people0 even educated people0
thou!ht it %asn(t1 #t didn(t have to e true0 ut it is1 We kno% this ecause a risin! flood of evidence
supports it1 Evolution is a fact0 and this ook %ill demonstrate it1 *o reputale scientist disputes it0
and no uniased reader %ill close the ook doutin! it1
Why0 then0 do %e speak of ('ar%in(s theory of evolution(0 therey0 it seems0 !ivin! spurious
comfort to those of a creationist persuasion : the history:deniers0 the >5:percenters : %ho think the
%ord (theory( is a concession0 handin! them some kind of !ift or victoryB
W)&T #S & T)EOR6B W)&T #S & ,&CTB
Only a theoryB -et(s look at %hat (theory( means1 The !xford En$lish Dictionary !ives t%o
meanin!s 7actually more0 ut these are the t%o that matter here81
Theory, Sense 1: & scheme or system of ideas or statements held as an explanation or
account of a !roup of facts or phenomenaJ a hypothesis that has een confirmed or estalished y
oservation or experiment0 and is propounded or accepted as accountin! for the kno%n factsJ a
statement of %hat are held to e the !eneral la%s0 principles0 or causes of somethin! kno%n or
Theory, Sense 2: & hypothesis proposed as an explanationJ hence0 a mere hypothesis0
speculation0 conHectureJ an idea or set of ideas aout somethin!J an individual vie% or notion1
Oviously the t%o meanin!s are Euite different from one another1 &nd the short ans%er to
my Euestion aout the theory of evolution is that the scientists are usin! Sense 20 %hile the
creationists are : perhaps mischievously0 perhaps sincerely : optin! for Sense 31 & !ood example of
Sense 2 is the )eliocentric Theory of the Solar System0 the theory that Earth and the other planets
orit the sun1 Evolution fits Sense 2 perfectly1 'ar%in(s theory of evolution is indeed a (scheme or
system of ideas or statements(1 #t does account for a massive (!roup of facts or phenomena(1 #t is (a
hypothesis that has een confirmed or estalished y oservation or experiment( and0 y !enerally
informed consent0 it is (a statement of %hat are held to e the !eneral la%s0 principles0 or causes of
somethin! kno%n or oserved(1 #t is certainly very far from (a mere hypothesis0 speculation0
conHecture(1 Scientists and creationists are understandin! the %ord (theory( in t%o very different
senses1 Evolution is a theory in the same sense as the heliocentric theory1 #n neither case should the
%ord (only( e used0 as in (only a theory(1
&s for the claim that evolution has never een (proved(0 proof is a notion that scientists have
een intimidated into mistrustin!1 #nfluential philosophers tell us %e can(t prove anythin! in
science1 "athematicians can prove thin!s : accordin! to one strict vie%0 they are the only people
%ho can : ut the est that scientists can do is fail to disprove thin!s %hile pointin! to ho% hard
they tried1 Even the undisputed theory that the moon is smaller than the sun cannot0 to the
satisfaction of a certain kind of philosopher0 e proved in the %ay that0 for example0 the
Pytha!orean Theorem can e proved1 But massive accretions of evidence support it so stron!ly that
to deny it the status of (fact( seems ridiculous to all ut pedants1 The same is true of evolution1
Evolution is a fact in the same sense as it is a fact that Paris is in the *orthern )emisphere1 Thou!h
lo!ic:choppers rule the to%n0L some theories are eyond sensile dout0 and %e call them facts1 The
more ener!etically and thorou!hly you try to disprove a theory0 if it survives the assault0 the more
closely it approaches %hat common sense happily calls a fact1
# could carry on usin! (Theory Sense 2( and (Theory Sense 3( ut numers are unmemorale1
# need sustitute %ords1 We already have a !ood %ord for (Theory Sense 3(1 #t is (hypothesis(1
Everyody understands that a hypothesis is a tentative idea a%aitin! confirmation 7or falsification80
and it is precisely this tentativeness that evolution has no% shed0 althou!h it %as still urdened %ith
it in 'ar%in(s time1 (Theory Sense 2( is harder1 #t %ould e nice simply to !o on usin! (theory(0 as
thou!h (Sense 3( didn(t exist1 #ndeed0 a !ood case could e made that Sense 3 sho"ldn't exist0
ecause it is confusin! and unnecessary0 !iven that %e have (hypothesis(1 $nfortunately Sense 3 of
(theory( is in common use and %e can(t y fiat an it1 # am therefore !oin! to take the considerale0
ut Hust for!ivale0 lierty of orro%in! from mathematics the %ord (theorem( for Sense 21 #t is
actually a mis:orro%in!0 as %e shall see0 ut # think the risk of confusion is out%ei!hed y the
enefits1 &s a !esture of appeasement to%ards affronted mathematicians0 # am !oin! to chan!e my
spellin! to (theorum(1 L ,irst0 let me explain the strict mathematical usa!e of theorem0 %hile at the
same time clarifyin! my earlier statement that0 strictly speakin!0 only mathematicians are licensed
to proe anythin! 7la%yers aren(t0 despite %ell:remunerated pretensions81
To a mathematician0 a proof is a lo!ical demonstration that a conclusion necessarily follo%s
from axioms that are assumed1 Pytha!oras( Theorem is necessarily true0 provided only that %e
assume Euclidean axioms0 such as the axiom that parallel strai!ht lines never meet1 6ou are %astin!
your time measurin! thousands of ri!ht:an!led trian!les0 tryin! to find one that falsifies Pytha!oras(
Theorem1 The Pytha!oreans proved it0 anyody can %ork throu!h the proof0 it(s Hust true and that(s
that1 "athematicians use the idea of proof to make a distinction et%een a (conHecture( and a
(theorem(0 %hich ears a superficial resemlance to the !ED(s distinction et%een the t%o senses of
(theory(1 & conHecture is a proposition that looks true ut has never een proved1 #t %ill ecome a
theorem %hen it has een proved1 & famous example is the Goldach ConHecture0 %hich states that
any even inte!er can e expressed as the sum of t%o primes1 "athematicians have failed to
disprove it for all even numers up to 455 thousand million million million0 and common sense
%ould happily call it Goldach(s ,act1 *evertheless it has never een proved0 despite lucrative
priFes ein! offered for the achievement0 and mathematicians ri!htly refuse to place it on the
pedestal reserved for theorems1 #f anyody ever finds a proof0 it %ill e promoted from Goldach(s
ConHecture to Goldach(s Theorem0 or maye C(s Theorem %here C is the clever mathematician
%ho finds the proof1
Carl Sa!an made sarcastic use of the Goldach ConHecture in his riposte to people %ho
claim to have een aducted y aliens1
Occasionally0 # !et a letter from someone %ho is in (contact( %ith extraterrestrials1 # am
invited to (ask them anythin!(1 &nd so over the years #(ve prepared a little list of Euestions1 The
extraterrestrials are very advanced0 rememer1 So # ask thin!s like0 (Please provide a short proof of
,ermat(s -ast Theorem(1 Or the Goldach ConHecture 1 1 1 # never !et an ans%er1 On the other hand0
if # ask somethin! like (Should %e e !oodB( # almost al%ays !et an ans%er1 &nythin! va!ue0
especially involvin! conventional moral Hud!ements0 these aliens are extremely happy to respond
to1 But on anythin! specific0 %here there is a chance to find out if they actually kno% anythin!
eyond %hat most humans kno%0 there is only silence1
,ermat(s -ast Theorem0 like the Goldach ConHecture0 is a proposition aout numers to
%hich noody has found an exception1 Provin! it has een a kind of holy !rail for mathematicians
ever since 2<4@0 %hen Pierre de ,ermat %rote in the mar!in of an old mathematics ook0 (# have a
truly marvellous proof 1 1 1 %hich this mar!in is too narro% to contain1( #t %as finally proved y the
En!lish mathematician &ndre% Wiles in 299=1 Before that0 some mathematicians think it should
have een called a conHecture1 Given the len!th and complication of Wiles(s successful proof0 and
his reliance on advanced t%entieth:century methods and kno%led!e0 most mathematicians think
,ermat %as 7honestly8 mistaken in his claim to have proved it1 # tell the story only to illustrate the
difference et%een a conHecture and a theorem1
&s # said0 # am !oin! to orro% the mathematicians( term (theorem(0 ut #(m spellin! it
(theorum( to differentiate it from a mathematical theorem1 & scientific theorum such as evolution or
heliocentrism is a theory that conforms to the Oxford dictionary(s (Sense 2(1
M#tN has een confirmed or estalished y oservation or experiment0 and is propounded or
accepted as accountin! for the kno%n factsJ Mit isN a statement of %hat are held to e the !eneral
la%s0 principles0 or causes of somethin! kno%n or oserved1
& scientific theorum has not een : cannot e : proved in the %ay a mathematical theorem is
proved1 But common sense treats it as a fact in the same sense as the (theory( that the Earth is round
and not flat is a fact0 and the theory that !reen plants otain ener!y from the sun is a fact1 &ll are
scientific theorumsA supported y massive Euantities of evidence0 accepted y all informed
oservers0 undisputed facts in the ordinary sense of the %ord1 &s %ith all facts0 if %e are !oin! to e
pedantic0 it is undenialy possile that our measurin! instruments0 and the sense or!ans %ith %hich
%e read them0 are the victims of a massive confidence trick1 &s Bertrand Russell said0 (We may all
have come into existence five minutes a!o0 provided %ith ready:made memories0 %ith holes in our
socks and hair that needed cuttin!1( Given the evidence no% availale0 for evolution to e anythin!
other than a fact %ould reEuire a similar confidence trick y the creator0 somethin! that fe% theists
%ould %ish to credit1
#t is time no% to examine the dictionary definition of a (fact(1 )ere is %hat the !ED has to
say 7a!ain there are several definitions0 ut this is the relevant one8A
Fat: Somethin! that has really occurred or is actually the caseJ somethin! certainly kno%n
to e of this characterJ hence0 a particular truth kno%n y actual oservation or authentic testimony0
as opposed to %hat is merely inferred0 or to a conHecture or fictionJ a datum of experience0 as
distin!uished from the conclusions that may e ased upon it1
*otice that0 like a theorum0 a fact in this sense doesn(t have the same ri!orous status as a
proved mathematical theorem0 %hich follo%s inescapaly from a set of assumed axioms1 "oreover0
(actual oservation or authentic testimony( can e horrily fallile0 and is over:rated in courts of
la%1 Psycholo!ical experiments have !iven us some stunnin! demonstrations0 %hich should %orry
any Hurist inclined to !ive superior %ei!ht to (eye:%itness( evidence1 & famous example %as
prepared y Professor 'aniel .1 Simons at the $niversity of #llinois1 )alf a doFen youn! people
standin! in a circle %ere filmed for 3= seconds tossin! a pair of asketalls to each other0 and %e0
the experimental suHects0 %atch the film1 The players %eave in and out of the circle and chan!e
places as they pass and ounce the alls0 so the scene is Euite actively complicated1 Before ein!
sho%n the film0 %e are told that %e have a task to perform0 to test our po%ers of oservation1 We
have to count the total numer of times alls are passed from person to person1 &t the end of the
test0 the counts are duly %ritten do%n0 ut : little does the audience kno% : this is not the real testK
&fter sho%in! the film and collectin! the counts0 the experimenter drops his omshell1
(&nd ho% many of you sa% the !orillaB( The maHority of the audience looks affledA lank1 The
experimenter then replays the film0 ut this time tells the audience to %atch in a relaxed fashion
%ithout tryin! to count anythin!1 &maFin!ly0 nine seconds into the film0 a man in a !orilla suit
strolls nonchalantly to the centre of the circle of players0 pauses to face the camera0 thumps his
chest as if in elli!erent contempt for eye:%itness evidence0 and then strolls off %ith the same
insouciance as efore 7see colour pa!e ;81 )e is there in full vie% for nine %hole seconds : more
than one:third of the film : and yet the maHority of the %itnesses never see him1 They %ould s%ear
an oath in a court of la% that no man in a !orilla suit %as present0 and they %ould s%ear that they
had een %atchin! %ith more than usually acute concentration for the %hole 3= seconds0 precisely
ecause they %ere countin! all:passes1 "any experiments alon! these lines have een performed0
%ith similar results0 and %ith similar reactions of stupefied diselief %hen the audience is finally
sho%n the truth1 Eye:%itness testimony0 (actual oservation(0 (a datum of experience( : all are0 or at
least can e0 hopelessly unreliale1 #t is0 of course0 exactly this unreliaility amon! oservers that
sta!e conHurors exploit %ith their techniEues of delierate distraction1
The dictionary definition of a fact mentions (act"al obseration or authentic testimony0 as
opposed to what is merely inferred( 7emphasis added81 The implied peHorative of that (merely( is a it
of a cheek1 Careful inference can e more reliale than (actual oservation(0 ho%ever stron!ly our
intuition protests at admittin! it1 # myself %as flaer!asted %hen # failed to see the Simons !orilla0
and frankly incredulous that it had really een there1 Sadder and %iser after my second vie%in! of
the film0 # shall never a!ain e tempted to !ive eye%itness testimony an automatic preference over
indirect scientific inference1 The !orilla film0 or somethin! like it0 should perhaps e sho%n to all
Huries efore they retire to consider their verdicts1 &ll Hud!es too1
&dmittedly0 inference has to e ased ultimately on oservation y our sense or!ans1 ,or
example0 %e use our eyes to oserve the printout from a '*& seEuencin! machine0 or from the
-ar!e )adron Collider1 But : all intuition to the contrary : direct oservation of an alle!ed event
7such as a murder8 as it actually happens is not necessarily more reliale than indirect oservation
of its conseEuences 7such as '*& in a loodstain8 fed into a %ell:constructed inference en!ine1
"istaken identity is more likely to arise from direct eye:%itness testimony than from indirect
inference derived from '*& evidence1 &nd0 y the %ay0 there is a distressin!ly lon! list of people
%ho have een %ron!ly convicted on eye:%itness testimony and suseEuently freed : sometimes
after many years : ecause of ne% evidence from '*&1 #n Texas alone0 thirty:five condemned
people have een exonerated since '*& evidence ecame admissile in court1 &nd that(s Hust the
ones %ho are still alive1 Given the !usto %ith %hich the State of Texas enforces the death penalty
7durin! his six years as Governor0 Geor!e W1 Bush si!ned a death %arrant once a fortni!ht on
avera!e80 %e have to assume that a sustantial numer of executed people %ould have een
exonerated if '*& evidence had een availale in time for them1
This ook %ill take inference seriously : not mere inference ut proper scientific inference :
and # shall sho% the irrefra!ale po%er of the inference that evolution is a fact1 Oviously0 the vast
maHority of evolutionary chan!e is invisile to direct eye:%itness oservation1 "ost of it happened
efore %e %ere orn0 and in any case it is usually too slo% to e seen durin! an individual(s
lifetime1 The same is true of the relentless pullin! apart of &frica and South &merica0 %hich occurs0
as %e shall see in Chapter 90 too slo%ly for us to notice1 With evolution0 as %ith continental drift0
inference after the event is all that is availale to us0 for the ovious reason that %e don(t exist until
after the event1 But do not for one nanosecond underestimate the po%er of such inference1 The slo%
driftin! apart of South &merica and &frica is no% an estalished fact in the ordinary lan!ua!e sense
of (fact(0 and so is our common ancestry %ith porcupines and pome!ranates1
We are like detectives %ho come on the scene after a crime has een committed1 The
murderer(s actions have vanished into the past1 The detective has no hope of %itnessin! the actual
crime %ith his o%n eyes1 #n any case0 the !orilla:suit experiment and others of its kind have tau!ht
us to mistrust our o%n eyes1 What the detective does have is traces that remain0 and there is a !reat
deal to trust there1 There are footprints0 fin!erprints 7and no%adays '*& fin!erprints too80
loodstains0 letters0 diaries1 The %orld is the %ay the %orld should e if this and this history0 ut not
that and that history0 led up to the present1
The distinction et%een the t%o dictionary meanin!s of (theory( is not an unrid!eale
chasm0 as many historical examples sho%1 #n the history of science0 theorums often start off as
(mere( hypotheses1 -ike the theory of continental drift0 an idea may even e!in its career mired in
ridicule0 efore pro!ressin! y painful steps to the status of a theorum or undisputed fact1 This is
not a philosophically difficult point1 The fact that some %idely held past eliefs have een
conclusively proved erroneous doesn(t mean %e have to fear that future evidence %ill al%ays sho%
our present eliefs to e %ron!1 )o% vulnerale our present eliefs are depends0 amon! other
thin!s0 on ho% stron! the evidence for them is1 People used to think the sun %as smaller than the
Earth0 ecause they had inadeEuate evidence1 *o% %e have evidence0 %hich %as not previously
availale0 that sho%s conclusively that it is much lar!er0 and %e can e totally confident that this
evidence %ill never0 ever e superseded1 This is not a temporary hypothesis that has so far survived
disproof1 Our present eliefs aout many thin!s may e disproved0 ut %e can %ith complete
confidence make a list of certain facts that %ill never e disproved1 Evolution and the heliocentric
theory %eren(t al%ays amon! them0 ut they are no%1
Biolo!ists often make a distinction et%een the fact of evolution 7all livin! thin!s are
cousins80 and the theory of %hat drives it 7they usually mean natural selection0 and they may
contrast it %ith rival theories such as -amarck(s theory of (use and disuse( and the (inheritance of
acEuired characteristics(81 But 'ar%in himself thou!ht of oth as theories in the tentative0
hypothetical0 conHectural sense1 This %as ecause0 in those days0 the availale evidence %as less
compellin! and it %as still possile for reputale scientists to dispute oth evolution and natural
selection1 *o%adays it is no lon!er possile to dispute the fact of evolution itself : it has !raduated
to ecome a theorum or oviously supported fact : ut it could still 7Hust8 e douted that natural
selection is its maHor drivin! force1
'ar%in explained in his autoio!raphy ho% in 2;4; he %as readin! "althus(s !n
Pop"lation (for amusement( 7under the influence0 "att Ridley suspects0 of his rother Erasmus(s
formidaly intelli!ent friend0 )arriet "artineau8 and received the inspiration for natural selectionA
()ere0 then # had at last !ot a theory y %hich to %ork1( ,or 'ar%in0 natural selection %as a
hypothesis0 %hich mi!ht have een ri!ht or mi!ht have een %ron!1 )e thou!ht the same of
evolution itself1 What %e no% call the fact of evolution %as0 in 2;4;0 a hypothesis for %hich
evidence needed to e collected1 By the time 'ar%in came to pulish !n the !ri$in of Species in
2;=90 he had amassed enou!h evidence to propel evolution itself0 thou!h still not natural selection0
a lon! %ay to%ards the status of fact1 #ndeed0 it %as this elevation from hypothesis to%ards fact that
occupied 'ar%in for most of his !reat ook1 The elevation has continued until0 today0 there is no
lon!er a dout in any serious mind0 and scientists speak0 at least informally0 of the fact of evolution1
&ll reputale iolo!ists !o on to a!ree that natural selection is one of its most important drivin!
forces0 althou!h : as some iolo!ists insist more than others : not the only one1 Even if it is not the
only one0 # have yet to meet a serious iolo!ist %ho can point to an alternative to natural selection
as a drivin! force of adaptie evolution : evolution to%ards positive improvement1
#n the rest of this ook0 # shall demonstrate that evolution is an inescapale fact0 and
celerate its astonishin! po%er0 simplicity and eauty1 Evolution is %ithin us0 around us0 et%een
us0 and its %orkin!s are emedded in the rocks of aeons past1 Given that0 in most cases0 %e don(t
live lon! enou!h to %atch evolution happenin! efore our eyes0 %e shall revisit the metaphor of the
detective comin! upon the scene of a crime after the event and makin! inferences1 The aids to
inference that lead scientists to the fact of evolution are far more numerous0 more convincin!0 more
incontrovertile0 than any eye:%itness reports that have ever een used0 in any court of la%0 in any
century0 to estalish !uilt in any crime1 Proof eyond reasonale doutB Reasonable doutB That is
the understatement of all time1
L *ot my favourite 6eats line0 ut apt in this case1
L ,or the sake of decorum G Pronounce it theorum1
W )6 did it take so lon! for a 'ar%in to arrive on the sceneB What delayed humanity(s
tumlin! to that luminously simple idea %hich seems0 on the face of it0 so much easier to !rasp than
the mathematical ideas !iven us y *e%ton t%o centuries earlier : or0 indeed0 y &rchimedes t%o
millennia earlierB "any ans%ers have een su!!ested1 Perhaps minds %ere co%ed y the sheer time
it must take for !reat chan!e to occur : y the mismatch et%een %hat %e no% call !eolo!ical deep
time and the lifespan and comprehension of the person tryin! to understand it1 Perhaps it %as
reli!ious indoctrination that held us ack1 Or perhaps it %as the dauntin! complexity of a livin!
or!an such as an eye0 frei!hted as it is %ith the e!uilin! illusion of desi!n y a master en!ineer1
Proaly all those played a role1 But Ernst "ayr0 !rand old man of the neo:'ar%inian synthesis0
%ho died in 355= at the a!e of 2550 repeatedly voiced a different suspicion1 ,or "ayr0 the culprit
%as the ancient philosophical doctrine of : to !ive it its modern name : essentialism1 The discovery
of evolution %as held ack y the dead hand of Plato1 L
T)E 'E&' )&*' O, P-&TO
,or Plato0 the (reality( that %e think %e see is Hust shado%s cast on the %all of our cave y
the flickerin! li!ht of the camp fire1 -ike other classical Greek thinkers0 Plato %as at heart a
!eometer1 Every trian!le dra%n in the sand is ut an imperfect shado% of the true essence of
trian!le1 The lines of the essential trian!le are pure Euclidean lines %ith len!th ut no readth0 lines
defined as infinitely narro% and as never meetin! %hen parallel1 The an!les of the essential trian!le
really do add up to exactly t%o ri!ht an!les0 not a picosecond of arc more or less1 This is not true of
a trian!le dra%n in the sandA ut the trian!le in the sand0 for Plato0 is ut an unstale shado% of the
ideal0 essential trian!le1
Biolo!y0 accordin! to "ayr0 is pla!ued y its o%n version of essentialism1 Biolo!ical
essentialism treats tapirs and raits0 pan!olins and dromedaries0 as thou!h they %ere trian!les0
rhomuses0 paraolas or dodecahedrons1 The raits that %e see are %an shado%s of the perfect
(idea( of rait0 the ideal0 essential0 Platonic rait0 han!in! some%here out in conceptual space
alon! %ith all the perfect forms of !eometry1 ,lesh:and:lood raits may vary0 ut their variations
are al%ays to e seen as fla%ed deviations from the ideal essence of rait1
)o% desperately unevolutionary that picture isK The Platonist re!ards any chan!e in raits
as a messy departure from the essential rait0 and there %ill al%ays e resistance to chan!e : as if
all real raits %ere tethered y an invisile elastic cord to the Essential Rait in the Sky1 The
evolutionary vie% of life is radically opposite1 'escendants can depart indefinitely from the
ancestral form0 and each departure ecomes a potential ancestor to future variants1 #ndeed0 &lfred
Russel Wallace0 independent co:discoverer %ith 'ar%in of evolution y natural selection0 actually
called his paper (On the tendency of varieties to depart indefinitely from the ori!inal type(1
#f there is a (standard rait(0 the accolade denotes no more than the centre of a ell:shaped
distriution of real0 scurryin!0 leapin!0 variale unnies1 &nd the distriution shifts %ith time1 &s
!enerations !o y0 there may !radually come a point0 not clearly defined0 %hen the norm of %hat
%e call raits %ill have departed so far as to deserve a different name1 There is no permanent
raitiness0 no essence of rait han!in! in the sky0 Hust populations of furry0 lon!:eared0
copropha!ous0 %hisker:t%itchin! individuals0 sho%in! a statistical distriution of variation in siFe0
shape0 colour and proclivities1 What used to e the lon!er:eared end of the old distriution may find
itself the centre of a ne% distriution later in !eolo!ical time1 Given a sufficiently lar!e numer of
!enerations0 there may e no overlap et%een ancestral and descendant distriutionsA the lon!est
ears amon! the ancestors may e shorter than the shortest ears amon! the descendants1 &ll is fluid0
as another Greek philosopher0 )eraclitus0 saidJ nothin! fixed1 &fter a hundred million years it may
e hard to elieve that the descendant animals ever had raits for ancestors1 6et in no !eneration
durin! the evolutionary process %as the predominant type in the population far from the modal type
in the previous !eneration or the follo%in! !eneration1 This %ay of thinkin! is %hat "ayr called
pop"lation thinkin$1 Population thinkin!0 for him0 %as the antithesis of essentialism1 &ccordin! to
"ayr0 the reason 'ar%in %as such an unconscionale time arrivin! on the scene %as that %e all :
%hether ecause of Greek influence or for some other reason : have essentialism urned into our
mental '*&1
,or the mind encased in Platonic linkers0 a rait is a rait is a rait1 To su!!est that
raitkind constitutes a kind of shiftin! cloud of statistical avera!es0 or that today(s typical rait
mi!ht e different from the typical rait of a million years a!o or the typical rait of a million
years hence0 seems to violate an internal taoo1 #ndeed0 psycholo!ists studyin! the development of
lan!ua!e tell us that children are natural essentialists1 "aye they have to e if they are to remain
sane %hile their developin! minds divide thin!s into discrete cate!ories each entitled to a uniEue
noun1 #t is no %onder that &dam(s first task0 in the Genesis myth0 %as to !ive all the animals names1
&nd it is no %onder0 in "ayr(s vie%0 that %e humans had to %ait for our 'ar%in until %ell
into the nineteenth century1 To dramatiFe ho% very anti:essentialist evolution is0 consider the
follo%in!1 On the (population:thinkin!( evolutionary vie%0 every animal is linked to every other
animal0 say rait to leopard0 y a chain of intermediates0 each so similar to the next that every link
could in principle mate %ith its nei!hours in the chain and produce fertile offsprin!1 6ou can(t
violate the essentialist taoo more comprehensively than that1 &nd it is not some va!ue thou!ht:
experiment confined to the ima!ination1 On the evolutionary vie%0 there really is a series of
intermediate animals connectin! a rait to a leopard0 every one of %hom lived and reathed0 every
one of %hom %ould have een placed in exactly the same species as its immediate nei!hours on
either side in the lon!0 slidin! continuum1 #ndeed0 every one of the series %as the child of its
nei!hour on one side and the parent of its nei!hour on the other1 6et the %hole series constitutes
a continuous rid!e from rait to leopard : althou!h0 as %e shall see later0 there never %as a
(raipard(1 There are similar rid!es from rait to %omat0 from leopard to loster0 from every
animal or plant to every other1 "aye you have reasoned for yourself %hy this startlin! result
follo%s necessarily from the evolutionary %orld:vie%0 ut let me spell it out any%ay1 #(ll call it the
hairpin thou!ht experiment1
Take a rait0 any female rait 7aritrarily stick to females0 for convenienceA it makes no
difference to the ar!ument81 Place her mother next to her1 *o% place the !randmother next to the
mother and so on ack in time0 ack0 ack0 ack throu!h the me!ayears0 a seemin!ly endless line of
female raits0 each one sand%iched et%een her dau!hter and her mother1 We %alk alon! the line
of raits0 ack%ards in time0 examinin! them carefully like an inspectin! !eneral1 &s %e pace the
line0 %e(ll eventually notice that the ancient raits %e are passin! are Hust a little it different from
the modern raits %e are used to1 But the rate of chan!e %ill e so slo% that %e shan(t notice the
trend from !eneration to !eneration0 Hust as %e can(t see the motion of the hour hand on our %atches
: and Hust as %e can(t see a child !ro%in!0 %e can only see later that she has ecome a teena!er0 and
later still an adult1 &n additional reason %hy %e don(t notice the chan!e in raits from one
!eneration to another is that0 in any one century0 the variation %ithin the current population %ill
normally e !reater than the variation et%een mothers and dau!hters1 So if %e try to discern the
movement of the (hour hand( y comparin! mothers %ith dau!hters0 or indeed !randmothers %ith
!randdau!hters0 such sli!ht differences as %e may see %ill e s%amped y the differences amon!
the raits( friends and relations !amollin! in the meado%s round aout1
*evertheless0 steadily and imperceptily0 as %e retreat throu!h time0 %e shall reach
ancestors that look less and less like a rait and more and more like a shre% 7and not very like
either81 One of these creatures #(ll call the hairpin end0 for reasons that %ill ecome apparent1 This
animal is the most recent common ancestor 7in the female line0 ut that is not important8 that raits
share %ith leopards1 We don(t kno% exactly %hat it looked like0 ut it follo%s from the evolutionary
vie% that it definitely had to exist1 -ike all animals0 it %as a memer of the same species as its
dau!hters and its mother1 We no% continue our %alk0 except that %e have turned the end in the
hairpin and are %alkin! for%ards in time0 aimin! to%ards the leopards 7amon! the hairpin(s many
and diverse descendants0 for %e shall continually meet forks in the line0 %here %e consistently
choose the fork that %ill eventually lead to leopards81 Each shre%:like animal alon! our for%ard
%alk is no% follo%ed y her dau!hter1 Slo%ly0 y imperceptile de!rees0 the shre%:like animals
%ill chan!e0 throu!h intermediates that mi!ht not resemle any modern animal much ut stron!ly
resemle each other0 perhaps passin! throu!h va!uely stoat:like intermediates0 until eventually0
%ithout ever noticin! an arupt chan!e of any kind0 %e arrive at a leopard1
+arious thin!s must e said aout this thou!ht experiment1 ,irst0 %e happen to have chosen
to %alk from rait to leopard0 ut # repeat that %e could have chosen porcupine to dolphin0 %allay
to !iraffe or human to haddock1 The point is that for any t%o animals there has to e a hairpin path
linkin! them0 for the simple reason that every species shares an ancestor %ith every other speciesA
all %e have to do is %alk ack%ards from one species to the shared ancestor0 then turn throu!h a
hairpin end and %alk for%ards to the other species1
Second0 notice that %e are talkin! only aout locatin! a chain of animals that links a modern
animal to another modern animal1 We are most emphatically not eolin$ a rait into a leopard1 #
suppose you could say %e are de0 evolvin! ack to the hairpin0 then evolvin! for%ards to the
leopard from there1 &s %e(ll see in a later chapter0 it is unfortunately necessary to explain0 a!ain and
a!ain0 that modern species don(t evolve into other modern species0 they Hust share ancestorsA they
are cousins1 This0 as %e shall see0 is also the ans%er to that disEuietin!ly common plaintA (#f humans
have evolved from chimpanFees0 ho% come there are still chimpanFees aroundB(
Third0 on our for%ard march from the hairpin animal0 %e aritrarily choose the path leadin!
to the leopard1 This is a real path of evolutionary history0 ut0 to repeat this important point0 %e
choose to i!nore numerous ranch points %here %e could have follo%ed evolution to countless
other end points : for the hairpin animal is the !rand ancestor not only of raits and leopards ut of
a lar!e fraction of modern mammals1
The fourth point0 %hich # have already emphasiFed0 is that0 ho%ever radical and extensive
the differences et%een the ends of the hairpin : rait and leopard0 say : each step alon! the chain
that links them is very0 very small1 Every individual alon! the chain is as similar to its nei!hours in
the chain as mothers and dau!hters are expected to e1 &nd more similar to its nei!hours in the
chain0 as # have also mentioned0 than to typical memers of the surroundin! population1
6ou can see ho% this thou!ht experiment drives a coach and horses throu!h the ele!ant
Greek temple of Platonic ideal forms1 &nd you can see ho%0 if "ayr is ri!ht that humans are deeply
imued %ith essentialist preconceptions0 he mi!ht %ell also e ri!ht aout %hy %e historically
found evolution so hard to stomach1
The %ord (essentialism( itself %asn(t invented till 29>= and so %as not availale to 'ar%in1
But he %as only too familiar %ith the iolo!ical version of it in the form of the (immutaility of
species(0 and much of his effort %as directed to%ards comatin! it under that name1 #ndeed0 in
several of 'ar%in(s ooks : more so in others than !n the !ri$in of Species itself : you(ll
understand fully %hat he(s on aout only if you shed modern presuppositions aout evolution0 and
rememer that a lar!e part of his audience %ould have een essentialists %ho never douted the
immutaility of species1 One of 'ar%in(s most tellin! %eapons in ar!uin! a!ainst this supposed
immutaility %as the evidence from domestication0 and it is domestication that %ill occupy the rest
of this chapter1
Sculptin! the !ene pool
'ar%in kne% plenty aout animal and plant reedin!1 )e communed %ith pi!eon fanciers
and horticulturalists0 and he loved do!s1L *ot only is the first chapter of !n the!ri$in of Species all
aout domestic varieties of animals and plantsJ 'ar%in also %rote a %hole ook on the suHect1 The
1ariation of Animals and Plants "nder Domestication has chapters on do!s and cats0 horses and
asses0 pi!s0 cattle0 sheep and !oats0 raits0 pi!eons 7t%o chaptersJ pi!eons %ere a particular love of
'ar%in80 chickens and various other irds0 and plants0 includin! the amaFin! caa!es1 Caa!es
are a ve!etale affront to essentialism and the immutaility of species1 The %ild caa!e0 Brassica
oleracea0 is an undistin!uished plant0 va!uely like a %eedy version of a domestic caa!e1 #n Hust a
fe% centuries0 %ieldin! the fine and coarse chisels furnished y the toolox of selective reedin!
techniEues0 horticulturalists have sculpted this rather nondescript plant into ve!etales as strikin!ly
different from each other and from the %ild ancestor as roccoli0 cauliflo%er0 kohlrai0 kale0
Brussels sprouts0 sprin! !reens0 romanescu and0 of course0 the various kinds of ve!etales that are
still commonly called caa!e1
&nother familiar example is the sculptin! of the %olf0 #anis l"p"s0 into the t%o hundred or
so reeds of do!0 #anis familiaris0 that are reco!niFed as separate y the $D Dennel Clu0 and the
lar!er numer of reeds that are !enetically isolated from one another y the apartheid:like rules of
pedi!ree reedin!1
#ncidentally0 the %ild ancestor of all domestic do!s really does seem to e the %olf and only
the %olf 7althou!h its domestication may have happened independently in different places around
the %orld81 Evolutionists haven(t al%ays thou!ht so1 'ar%in0 alon! %ith many of his
contemporaries0 suspected that several species of %ild canid0 includin! %olves and Hackals0 had
contriuted ancestry to our domestic do!s1 The *oel PriFe:%innin! &ustrian etholo!ist Donrad
-orenF %as of the same vie%1 )is %an %eets Do$0 pulished in 29>90 pushes the notion that
domestic do! reeds fall into t%o main !roupsA those derived from Hackals 7the maHority8 and those
derived from %olves 7-orenF(s o%n favourites0 includin! Cho%s81 -orenF seems to have had no
evidence at all for his dichotomy0 other than the differences that he thou!ht he sa% in the
personalities and characters of the reeds1 The matter remained open until molecular !enetic
evidence came alon! to clinch it1 There is no% no dout1 'omestic do!s have no Hackal ancestry at
all1 &ll reeds of do!s are modified %olvesA not Hackals0 not coyotes and not foxes1
The main point # %ant to dra% out of domestication is its astonishin! po%er to chan!e the
shape and ehaviour of %ild animals0 and the speed %ith %hich it does so1 Breeders are almost like
modellers %ith endlessly malleale clay0 or like sculptors %ieldin! chisels0 carvin! do!s or horses0
or co%s or caa!es0 to their %him1 # shall return to this ima!e shortly1 The relevance to natural
evolution is that0 althou!h the selectin! a!ent is man and not nature0 the process is other%ise exactly
the same1 This is %hy 'ar%in !ave so much prominence to domestication at the e!innin! of !n
the !ri$in of Species1 &nyody can understand the principle of evolution y artificial selection1
*atural selection is the same0 %ith one minor detail chan!ed1
Strictly speakin!0 it is not the ody of the do! or the caa!e that is carved y the
reederGsculptor ut the !ene pool of the reed or species1 The idea of a !ene pool is central to the
ody of kno%led!e and theory that !oes under the name of the (*eo:'ar%inian Synthesis(1 'ar%in
himself kne% nothin! of it1 #t %as not a part of his intellectual %orld0 nor indeed %ere !enes1 )e
%as a%are0 of course0 that characteristics run in familiesJ a%are that offsprin! tend to resemle their
parents and silin!sJ a%are that particular characteristics of do!s and pi!eons reed true1 )eredity
%as a central plank of his theory of natural selection1 But a !ene pool is somethin! else1 The
concept of a !ene pool has meanin! only in the li!ht of "endel(s la% of the independent assortment
of hereditary particles1 'ar%in never kne% "endel(s la%s0 for althou!h Gre!or "endel0 the
&ustrian monk %ho %as the father of !enetics0 %as 'ar%in(s contemporary0 he pulished his
findin!s in a German Hournal %hich 'ar%in never sa%1
& "endelian !ene is an all:or:nothin! entity1 When you %ere conceived0 %hat you received
from your father %as not a sustance0 to e mixed %ith %hat you received from your mother as if
mixin! lue paint and red paint to make purple1 #f this %ere really ho% heredity %orked 7as people
va!uely thou!ht in 'ar%in(s time8 %e(d all e a middlin! avera!e0 half%ay et%een our t%o parents1
#n that case0 all variation %ould rapidly disappear from the population 7no matter ho% assiduously
you mix purple paint %ith purple paint0 you(ll never reconstitute the ori!inal red and lue81 #n fact0
of course0 anyody can plainly see that there is no such intrinsic tendency for variation to decrease
in a population1 "endel sho%ed that this is ecause %hen paternal !enes and maternal !enes are
comined in a child 7he didn(t use the %ord (!ene(0 %hich %asn(t coined until 295980 it is not like
lendin! paints0 it is more like shufflin! and reshufflin! cards in a pack1 *o%adays0 %e kno% that
!enes are len!ths of '*& code0 not physically separate like cards0 ut the principle remains valid1
Genes don(t lendJ they shuffle1 6ou could say they are shuffled adly0 %ith !roups of cards
stickin! to!ether for several !enerations of shufflin! efore chance happens to split them1
&ny one of your e!!s 7or sperms if you are male8 contains either your father(s version of a
particular !ene or your mother(s version0 not a lend of the t%o1 &nd that particular !ene came from
one and only one of your four !randparentsJ and from one and only one of your ei!ht !reat:
!randparents1 L
)indsi!ht says this should have een ovious all alon!1 When you cross a male %ith a
female0 you expect to !et a son or a dau!hter0 not a hermaphrodite1O )indsi!ht says anyody in an
armchair could have !eneraliFed the same all:or:none principle to the inheritance of each and every
characteristic1 ,ascinatin!ly0 'ar%in himself %as !limmerin!ly close to this0 ut he stopped Hust
short of makin! the full connection1 #n 2;<< he %rote0 in a letter to &lfred WallaceA
"y dear Wallace
# do not think you understand %hat # mean y the non:lendin! of certain varieties1 #t does
not refer to fertility1 &n instance %ill explain1 # crossed the Painted -ady and Purple s%eet peas0
%hich are very differently coloured varieties0 and !ot0 even out of the same pod0 oth varieties
perfect ut none intermediate1 Somethin! of this kind0 # should think0 must occur at first %ith your
utterflies 1 1 1 Thou!h these cases are in appearance so %onderful0 # do not kno% that they are really
more so than every female in the %orld producin! distinct male and female offsprin!1
'ar%in came that close to discoverin! "endel(s la% of the non:lendin! of 7%hat %e %ould
no% call8 !enes1 L The case is analo!ous to the claim0 y various a!!rieved apolo!ists0 that other
+ictorian scientists0 for example Patrick "atthe% and Ed%ard Blyth0 had discovered natural
selection efore 'ar%in did1 #n a sense that is true0 as 'ar%in ackno%led!ed0 ut # think the
evidence sho%s that they didn(t understand ho% important it is1 $nlike 'ar%in and Wallace0 they
didn(t see it as a $eneral phenomenon %ith universal si!nificance : %ith the po%er to drive the
evolution of all livin! thin!s in the direction of positive improvement1 #n the same %ay0 this letter to
Wallace sho%s that 'ar%in !ot tantaliFin!ly close to !raspin! the point aout the non:lendin!
nature of heredity1 But he didn(t see its !enerality0 and in particular he failed to see it as the ans%er
to the riddle of %hy variation didn(t automatically disappear from populations1 That %as left to
t%entieth:century scientists0 uildin! on "endel(s efore:his:time discovery1 O
So no% the concept of the !ene pool starts to make sense1 & sexually reproducin!
population0 such as0 say0 all the rats on &scension #sland0 remotely isolated in the South &tlantic0 is
continually shufflin! all the !enes on the island1 There is no intrinsic tendency for each !eneration
to ecome less variale than the previous !eneration0 no tendency to%ards ever more orin!ly !rey0
middlin! intermediates1 The !enes remain intact0 shuffled aout from individual ody to individual
ody as the !enerations !o y0 ut not blendin$ %ith one another0 never contaminatin! each other1
&t any one time0 the !enes are all sittin! in the odies of individual rats0 or they are movin! into
ne% rat odies via sperms1 But if %e take a lon! vie% across many !enerations0 %e see all the rat
!enes on the island ein! mixed up as thou!h they %ere cards in a sin!le %ell:shuffled packA one
sin!le pool of !enes1
#(m !uessin! that the rat !ene pool on a small and isolated island such as &scension is a self:
contained and rather %ell:stirred pool0 in the sense that the recent ancestors of any one rat could
have lived any%here on the island0 ut proaly not any%here other than on the island0 !ive or take
the occasional sto%a%ay on a ship1 But the !ene pool of the rats on a lar!e land mass such as
Eurasia %ould e much more complicated1 & rat livin! in "adrid %ould derive most of its !enes
from ancestors livin! in the %estern end of the Eurasian continent rather than0 say0 "on!olia or
Sieria0 not ecause of specific arriers to !ene flo% 7thou!h those exist too8 ut ecause of the
sheer distances involved1 #t takes time for sexual shufflin! to %ork a !ene from one side of a
continent to the other1 Even if there are no physical arriers such as rivers or mountain ran!es0 !ene
flo% across such a lar!e land mass %ill still e slo% enou!h for the !ene pool to deserve the name
(viscous(1 & rat livin! in +ladivostok %ould trace most of its !enes ack to ancestors in the east1 The
Eurasian !ene pool %ould e shuffled0 as on &scension #sland0 ut not homo!eneously shuffled
ecause of the distances involved1 "oreover0 partial arriers such as mountain ran!es0 lar!e rivers
or deserts %ould further !et in the %ay of homo!eneous shufflin!0 therey structurin! and
complicatin! the !ene pool1 These complications don(t devalue the idea of the !ene pool1 The
perfectly stirred !ene pool is a useful astraction0 like a mathematician(s astraction of a perfect
strai!ht line1 Real !ene pools0 even on small islands like &scension0 are imperfect approximations0
only partially shuffled1 The smaller and less roken:up the island0 the etter the approximation to
the astract ideal of the perfectly stirred !ene pool1
.ust to round off the thou!ht aout !ene pools0 each individual animal that %e see in a
population is a samplin$ of the !ene pool of its time 7or rather its parents( time81 There is no
intrinsic tendency in !ene pools for particular !enes to increase or decrease in freEuency1 But %hen
there is a systematic increase or decrease in the freEuency %ith %hich %e see a particular !ene in a
!ene pool0 that is precisely and exactly %hat is meant y evolution1 The Euestion0 therefore0
ecomesA why should there e a systematic increase or decrease in a !ene(s freEuencyB That0 of
course0 is %here thin!s start to !et interestin!0 and %e shall come to it in due course1
Somethin! funny happens to the !ene pools of domestic do!s1 Breeders of pedi!ree
Pekineses or 'almatians !o to elaorate len!ths to stop !enes crossin! from one !ene pool to
another1 Stud ooks are kept0 !oin! ack many !enerations0 and misce!enation is the %orst thin!
that can happen in the ook of a pedi!ree reeder1 #t is as thou!h each reed of do! %ere
incarcerated on its o%n little &scension #sland0 kept apart from every other reed1 But the arrier to
interreedin! is not lue %ater ut human rules1 Geo!raphically the reeds all overlap0 ut they
mi!ht as %ell e on separate islands ecause of the %ay their o%ners police their matin!
opportunities1 Of course0 from time to time the rules are roken1 -ike a rat sto%in! a%ay on a ship
to &scension #sland0 a %hippet itch0 say0 escapes the leash and mates %ith a spaniel1 But the
mon!rel puppies that result0 ho%ever loved they may e as individuals0 are cast off the island
laelled Pedi!ree Whippet1 The island itself remains a pure %hippet island1 Other pure:red
%hippets ensure that the !ene pool of the virtual island laelled Whippet continues uncontaminated1
There are hundreds of man:made (islands(0 one for each reed of pedi!ree do!1 Each one is a virtual
island0 in the sense that it is not !eo!raphically localiFed1 Pedi!ree %hippets or Pomeranians are to
e found in many different places around the %orld0 and cars0 ships and planes are used to ferry the
!enes from one !eo!raphical place to another1 The virtual !enetic island that is the Pekinese !ene
pool overlaps !eo!raphically0 ut not !enetically 7except %hen a itch reaks cover80 %ith the
virtual !enetic island that is the oxer !ene pool and the virtual island that is the St Bernard !ene
*o% let(s return to the remark that opened my discussion of !ene pools1 # said that if human
reeders are to e seen as sculptors0 %hat they are carvin! %ith their chisels is not do! flesh ut
!ene pools1 #t appears to e do! flesh ecause the reeder mi!ht announce an intention to0 say0
shorten the snouts of future !enerations of oxers1 &nd the end product of such an intention %ould
indeed e a shorter snout0 as thou!h a chisel had een taken to the ancestor(s face1 But0 as %e have
seen0 a typical oxer in any one !eneration is a samplin! of the contemporary !ene pool1 #t is the
!ene pool that has een carved and %hittled over the years1 Genes for lon! snouts have een
chiselled out of the !ene pool and replaced y !enes for short snouts1 Every reed of do!0 from
dachshund to 'almatian0 from oxer to orFoi0 from poodle to Pekinese0 from Great 'ane to
chihuahua0 has een carved0 chiselled0 kneaded0 moulded0 not literally as flesh and one ut in its
!ene pool1
#t isn(t all done y carvin!1 "any of our familiar reeds of do! %ere ori!inally derived as
hyrids of other reeds0 often Euite recently0 for example in the nineteenth century1 )yridiFation0
of course0 represents a delierate violation of the isolation of the !ene pools on virtual islands1
Some hyridiFation schemes are desi!ned %ith such care that the reeders %ould resent their
products ein! descried as mon!rels or mutts 7as President Oama deli!htfully descried himself81
The (-aradoodle( is a hyrid et%een a standard poodle and a -arador retriever0 the result of a
carefully crafted Euest for the est virtues of oth reeds1 -aradoodle o%ners have estalished
societies and associations Hust like those of pure:red pedi!ree do!s1 There are t%o schools of
thou!ht in the -aradoodle ,ancy0 and those of other such desi!ner hyrids1 There are those %ho
are happy to !o on makin! -aradoodles y matin! poodles and -aradors to!ether1 &nd there are
those %ho are tryin! to initiate a ne% -aradoodle !ene pool that %ill reed true0 %hen
-aradoodles are mated to!ether1 &t present0 second:!eneration -aradoodle !enes recomine to
produce more variety than pure:red pedi!ree do!s are supposed to sho%1 This is ho% many (pure(
reeds !ot their startA they %ent throu!h an intermediate sta!e of hi!h variation0 suseEuently
trimmed do%n throu!h !enerations of careful reedin!1
Sometimes0 ne% reeds of do! !et their start %ith the adoption of a sin!le maHor mutation1
"utations are the random chan!es in !enes that constitute the ra% material for evolution y non:
random selection1 #n nature0 lar!e mutations seldom survive0 ut !eneticists like them in the
laoratory ecause they are easy to study1 Breeds of do! %ith very short le!s0 like asset hounds
and dachshunds0 acEuired them in a sin!le step %ith the !enetic mutation called achondroplasia0 a
classic example of a lar!e mutation that %ould e unlikely to survive in nature1 & similar mutation
is responsile for the commonest kind of human d%arfismA the trunk is of nearly normal siFe0 ut
the le!s and arms are short1 Other !enetic routes produce miniature reeds that retain the
proportions of the ori!inal1 'o! reeders can achieve chan!es in siFe and shape y selectin!
cominations of a fe% maHor mutations such as achondroplasia and lots of minor !enes1 *or do they
need to understand the !enetics in order to achieve chan!e effectively1 Without any understandin!
at all0 Hust y choosin! %ho mates %ith %hom0 you can reed for all kinds of desired characteristics1
This is %hat do! reeders0 and animal and plant reeders !enerally0 achieved for centuries efore
anyody understood anythin! aout !enetics1 &nd there(s a lesson in that aout natural selection0 for
nature0 of course0 has no understandin! or a%areness of anythin! at all1
The &merican Foolo!ist Raymond Coppin!er makes the point that puppies of different
reeds are much more similar to each other than adult do!s are1 Puppies can(t afford to e different0
ecause the main thin! they have to do is suck0L and suckin! presents pretty much the same
challen!es for all reeds1 #n particular0 in order to e !ood at suckin!0 a puppy can(t have a lon!
snout like a orFoi or a retriever1 That(s %hy all puppies look like pu!s1 6ou could say that an adult
pu! is a puppy %hose face didn(t properly !ro% up1 "ost do!s0 after they are %eaned0 develop a
relatively lon!er snout1 Pu!s0 ulldo!s and Pekineses don(tJ they !ro% in other departments0 %hile
the snout retains its infantile proportions1 The technical term for this is neoteny0 and %e(ll meet it
a!ain %hen %e come on to human evolution in Chapter @1
#f an animal !ro%s at the same rate in all its parts0 so that the adult is Hust a uniformly
inflated replica of the infant0 it is said to !ro% isometrically1 #sometric !ro%th is Euite rare1 #n
allometric !ro%th0 y contrast0 different parts !ro% at different rates1 Often0 the rates of !ro%th of
different parts of an animal ear some simple mathematical relation to each other0 a phenomenon
that %as investi!ated especially y Sir .ulian )uxley in the 2945s1 'ifferent reeds of do! achieve
their different shapes y means of !enes that chan!e the allometric !ro%th relationships et%een
the parts of the ody1 ,or example0 ulldo!s !et their Churchillian sco%l from a !enetic tendency
to%ards slo%er !ro%th of the nasal ones1 This has knock:on effects on the relative !ro%th of the
surroundin! ones0 and indeed all the surroundin! tissues1 One of these knock:on effects is that the
palate is pulled up into an a%k%ard position0 so the ulldo!(s teeth stick out and it has a tendency to
drile1 Bulldo!s also have reathin! difficulties0 %hich are shared y Pekineses1 Bulldo!s even
have difficulty ein! orn ecause the head is disproportionately i!1 "ost if not all the ulldo!s
you see today %ere orn y caesarian section1
BorFois are the opposite1 They have extra lon! snouts1 #ndeed0 they are unusual in that the
elon!ation of the snout e!ins efore they are orn0 %hich proaly makes orFoi puppies less
proficient suckers than other reeds1 Coppin!er speculates that the human desire to reed orFois
for lon! snouts has reached a limit imposed y the survival capacity of puppies tryin! to suck1
What lessons do %e learn from the domestication of the do!B ,irst0 the !reat variety amon!
reeds of do!s0 from Great 'anes to 6orkies0 from Scotties to &iredales0 from rid!eacks to
dachshunds0 from %hippets to St Bernards0 demonstrates ho% easy it is for the non:random
selection of !enes : the (carvin! and %hittlin!( of !ene pools : to produce truly dramatic chan!es in
anatomy and ehaviour0 and so fast1 Surprisin!ly fe% !enes may e involved1 6et the chan!es are
so lar!e : the differences et%een reeds so dramatic : that you mi!ht expect their evolution to take
millions of years instead of Hust a matter of centuries1 #f so much evolutionary chan!e can e
achieved in Hust a fe% centuries or even decades0 Hust think %hat mi!ht e achieved in ten or a
hundred million years1
+ie%in! the process over centuries0 it is no empty fancy that human do! reeders have
seiFed do! flesh like modellin! clay and pushed it0 pulled it0 kneaded it into shape0 more or less at
%ill1 Of course0 as # pointed out earlier0 %e have really een kneadin! not do! flesh ut do! !ene
pools1 &nd (carved( is a etter metaphor than (kneaded(1 Some sculptors %ork y takin! a lump of
clay and kneadin! it into shape1 Others take a lump of stone or %ood0 and carve it y s"btractin$
its %ith a chisel1 Oviously do! fanciers don(t carve do!s into shape y sutractin! its of do!
flesh1 But they do somethin! close to carvin! do! !ene pools y sutraction1 #t is more complicated
than pure sutraction0 ho%ever1 "ichelan!elo took a sin!le chunk of marle0 and then sutracted
marle from it to reveal 'avid lurkin! inside1 *othin! %as added1 Gene pools0 on the other hand0
are continually added to0 for example y mutation0 %hile at the same time non:random death
sutracts1 The analo!y to sculpture reaks do%n here0 and should not e pushed too tenaciously0 as
%e(ll see a!ain in Chapter ;1
The idea of sculpture calls to mind the over:muscled physiEues of human ody:uilders0 and
non:human eEuivalents such as the Bel!ian Blue reed of cattle1 This %alkin! eef factory has een
contrived via a particular !enetic alteration called (doule musclin!(1 There is a sustance called
myostatin0 %hich limits muscle !ro%th1 #f the !ene that makes myostatin is disaled0 muscles !ro%
lar!er than usual1 #t is Euite often the case that a !iven !ene can mutate in more than one %ay to
produce the same outcome0 and indeed there are various %ays in %hich the myostatin:producin!
!ene can e disaled0 %ith the same effect1 &nother example is the reed of pi! called the Black
Exotic0 and there are individual do!s of various reeds that sho% the same exa!!erated musculature
for the same reason1 )uman ody:uilders achieve a similar physiEue y an extreme re!ime of
exercise0 and often y the use of anaolic steroidsA oth environmental manipulations that mimic
the !enes of the Bel!ian Blue and the Black Exotic1 The end result is the same0 and that is a lesson
in itself1 Genetic and environmental chan!es can produce identical outcomes1 #f you %anted to rear
a human child to %in a ody:uildin! contest and you had a fe% centuries to spare0 you could start
y !enetic manipulation0 en!ineerin! exactly the same freak !ene as characteriFes Bel!ian Blue
cattle and Black Exotic pi!s1 #ndeed0 there are some humans kno%n to have deletions of the
myostatin !ene0 and they tend to e anormally %ell muscled1 #f you started %ith a mutant child and
made it pump iron as %ell 7presumaly the cattle and pi!s could not e caHoled into this80 you could
proaly end up %ith somethin! more !rotesEue than "r $niverse1
Political opposition to eu!enic reedin! of humans sometimes spills over into the almost
certainly false assertion that it is impossile1 *ot only is it immoral0 you may hear it said0 it
%ouldn(t %ork1 $nfortunately0 to say that somethin! is morally %ron!0 or politically undesirale0 is
not to say that it %ouldn(t %ork1 # have no dout that0 if you set your mind to it and had enou!h time
and enou!h political po%er0 you could reed a race of superior ody:uilders0 or hi!h:Humpers0 or
shot:puttersJ pearl fishers0 sumo %restlers0 or sprintersJ or 7# suspect0 althou!h no% %ith less
confidence ecause there are no animal precedents8 superior musicians0 poets0 mathematicians or
%ine:tasters1 The reason # am confident aout selective reedin! for athletic pro%ess is that the
Eualities needed are so similar to those that demonstraly %ork in the reedin! of racehorses and
carthorses0 of !reyhounds and sled!e do!s1 The reason # am still pretty confident aout the practical
feasiility 7thou!h not the moral or political desiraility8 of selective reedin! for mental or
other%ise uniEuely human traits is that there are so fe% examples %here an attempt at selective
reedin! in animals has ever failed0 even for traits that mi!ht have een thou!ht surprisin!1 Who
%ould have thou!ht0 for example0 that do!s could e red for sheep:herdin! skills0 or (pointin!(0 or
6ou %ant hi!h milk yield in co%s0 orders of ma!nitude more !allons than could ever e
needed y a mother to rear her aiesB Selective reedin! can !ive it to you1 Co%s can e modified
to !ro% vast and un!ainly udders0 and these continue to yield copious Euantities of milk
indefinitely0 lon! after the normal %eanin! period of a calf1 &s it happens0 dairy horses have not
een red in this %ay0 ut %ill anyone contest my et that %e could do it if %e triedB &nd of course0
the same %ould e true of dairy humans0 if anyone %anted to try1 &ll too many %omen0
amooFled y the myth that reasts like melons are attractive0 pay sur!eons lar!e sums of money
to implant silicone0 %ith 7for my money8 unappealin! results1 'oes anyone dout that0 !iven
enou!h !enerations0 the same deformity could e achieved y selective reedin!0 after the manner
of ,riesian co%sB
&out t%enty:five years a!o # developed a computer simulation to illustrate the po%er of
artificial selectionA a kind of computer !ame eEuivalent to reedin! priFe roses or do!s or cattle1
The player is faced %ith an array of nine shapes on the screen : (computer iomorphs( : the middle
one of %hich is the (parent( of the surroundin! ei!ht1 &ll the shapes are constructed under the
influence of a doFen or so (!enes(0 %hich are simply numers handed do%n from (parent( to
(offsprin!(0 %ith the possiility of small (mutations( intervenin! on the %ay1 & mutation is Hust a
sli!ht increment or decrement in the numerical value of the parent(s !ene1 Each shape is constructed
under the influence of a particular set of numers0 %hich are its o%n particular values of the doFen
!enes1 The player looks over the array of nine shapes and sees no !enes ut chooses the preferred
(ody( shape she %ants to reed from1 The other ei!ht iomorphs disappear from the screen0 the
chosen one !lides to the centre0 and (spa%ns( ei!ht ne% mutant (children(1 The process repeats for as
many (!enerations( as the player has time for0 and the avera!e shape of the (or!anisms( on the screen
!radually (evolves( as the !enerations !o y1 Only !enes are passed from !eneration to !eneration0
so0 y directly choosin! iomorphs y eye0 the player is inadvertently choosin! !enes1 That is Hust
%hat happens %hen reeders choose do!s or roses to reed from1
!io"or#hs $ro" the '!lin% Wath"a&er' #ro'ra"
So much for the !enetics1 The !ame starts to !et interestin! %hen %e consider the
(emryolo!y(1 The emryolo!y of a iomorph on the screen is the process y %hich its (!enes( :
those numerical values : influence its shape1 "any very different emryolo!ies can e ima!ined0
and # have tried out Euite a fe% of them1 "y first pro!ram0 called (Blind Watchmaker(0 uses a tree:
!ro%in! emryolo!y1 & main (trunk( sprouts t%o (ranches(0 then each ranch sprouts t%o ranches
of its o%n0 and so on1 The numer of ranches0 and their an!les and len!ths0 are all under !enetic
control0 determined y the numerical values of the !enes1 &n important feature of the ranchin! tree
emryolo!y is that it is rec"rsie1 # %on(t expound that idea here0 ut it means that a sin!le mutation
typically has an effect all over the tree0 rather than Hust in one corner of it1
&lthou!h the Blind Watchmaker pro!ram starts off %ith a simple ranchin! tree0 it rapidly
%anders off into a %onderland of evolved forms0 many %ith a stran!e eauty0 and some : dependin!
on the intentions of the human player : comin! to resemle familiar creatures such as insects0
spiders or starfish1 On the left is a (safari park( of creatures that Hust one player of the !ame 7me8
found in the y%ays and ack%aters of this stran!e computer %onderland1 #n a later version of the
pro!ram0 # expanded the emryolo!y to allo% for !enes controllin! the colour and shape of the
(ranches( of the tree1
& more elaorate pro!ram0 called (&rthromorphs(0 %hich # %rote Hointly %ith Ted Daehler0
then %orkin! for the &pple Computer Company0 emodies an (emryolo!y( %ith some interestin!
iolo!ical features specifically !eared to reedin! (insects(0 (spiders(0 (centipedes( and other creatures
resemlin! arthropods1 # have explained the arthromorphs in detail0 alon! %ith the iomorphs0
(conchomorphs( 7computer molluscs8 and other pro!rams in this vein0 in #limbin$ %o"nt
&s it happens0 the mathematics of shell emryolo!y are %ell understood0 so artificial
selection usin! my (conchomorph( pro!ram is capale of !eneratin! extremely lifelike forms 7see
over81 # shall refer ack to these pro!rams0 to make a completely different point0 in the final chapter1
)ere # have introduced them for the purpose of illustratin! the po%er of artificial selection0 even in
an extremely over:simplified computer environment1 #n the real %orld of a!riculture and
horticulture0 the %orld of the pi!eon fancier or do! reeder0 artificial selection can achieve so much
more1 Biomorphs0 arthromorphs and conchomorphs Hust illustrate the principle0 in somethin! like
the same %ay that artificial selection itself is !oin! to illustrate the principle ehind natural
selection : in the next chapter1
Conho"or#hs: o"#(ter)'enerate% shells sha#e% *y arti$iial seletion
'ar%in had first:hand experience of the po%er of artificial selection and he !ave it pride of
place in Chapter 2 of !n the !ri$in of Species1 )e %as softenin! his readers up to take delivery of
his o%n !reat insi!ht0 the po%er of natural selection1 #f human reeders can transform a %olf into a
Pekinese0 or a %ild caa!e into a cauliflo%er0 in Hust a fe% centuries or millennia0 %hy shouldn(t
the non:random survival of %ild animals and plants do the same thin! over millions of yearsB That
%ill e the conclusion of my next chapterJ ut my strate!y first %ill e to continue the softenin!:up
process0 to ease the passa!e to%ards understandin! of natural selection1
L This isn(t "ayr(s phrase0 thou!h it expresses his idea1
L Who could not love do!s0 they are such !ood sportsB
L This %ould e strictly true on the model of !enetics that "endel offered us0 and the model
of !enetics that all iolo!ists follo%ed until the Watson:Crick revolution of the 29=5s1 #t is nearly
ut not Euite true0 !iven %hat %e no% kno% aout !enes as lon! stretches of '*&1 ,or all practical
purposes %e can take it as true1
O On the farm %here # spent my childhood0 %e had one especially ostreperous and
a!!ressive co% called &rusha1 &rusha %as (a character( and a prolem1 One day the herdsman0 "r
Evans0 ruefully remarkedA (Seems to me0 &rusha is more like a cross et%een a ull and a co%1(
L There is a persistent0 ut false0 rumour that 'ar%in possessed a ound copy of the German
Hournal in %hich "endel pulished his results ut that the relevant pa!es %ere found uncut on
'ar%in(s death1 The meme proaly ori!inates from the fact that he possessed a ook called Die
Pflan2en0mischlin$e y W1 O1 ,ocke1 ,ocke did riefly refer to "endel0 and the pa!e %here he did
so %as indeed uncut in 'ar%in(s copy1 But ,ocke laid no special emphasis on "endel(s %ork and
sho%ed no evidence of understandin! its profound si!nificance0 so it is not ovious that 'ar%in
%ould have picked it out even if he had cut the relevant pa!e1 #n any case0 'ar%in(s German %as
not !reat1 #f he had read "endel(s paper0 the history of iolo!y %ould have een very different1 #t is
ar!uale that even "endel himself did not understand the full importance of his findin!s1 #f he had0
he mi!ht have %ritten to 'ar%in1 #n the lirary of "endel(s monastery in Brno0 # have held in my
hand "endel(s o%n copy 7in German8 of !n the !ri$in of Species and seen his mar!inalia0 %hich
indicate that he read it1
O Be!innin! in 295; %ith the endearin!ly eccentric0 cricket:lovin! mathematician G1 )1
)ardy and0 independently0 the German doctor Wilhelm Weiner!0 the theory culminated in the
%ork of the !reat !eneticist and statistician Ronald ,isher0 and0 a!ain lar!ely independently0 his co:
founders of population !enetics0 .1 B1 S1 )aldane and Se%all Wri!ht1
L *ot suckleA mothers suckle0 aies suck1
C)&PTER 3 sho%ed ho% the human eye0 %orkin! y selective reedin! over many
!enerations0 sculpted and kneaded do! flesh to assume a e%ilderin! variety of forms0 colours0
siFes and ehaviour patterns1 But %e are humans0 accustomed to makin! choices that are delierate
and planned1 &re there other animals that do the same thin! as human reeders0 perhaps %ithout
delieration or intention ut %ith similar resultsB 6es0 and they carry this ook(s softenin!:up
pro!ram steadily for%ard1 This chapter emarks on a step:y:step seduction of the mind as %e pass
from the familiar territory of do! reedin! and artificial selection to 'ar%in(s !iant discovery of
natural selection0 via some colourful intermediate sta!es1 The first of these intermediate steps alon!
the path of seduction 7is it over the top to call it a primrose pathB8 takes us into the honeyed %orld
of flo%ers1
Wild roses are a!reeale little flo%ers0 pretty enou!h0 ut nothin! to %rite home aout in the
terms one mi!ht lavish on0 say0 (Peace( or (-ovely -ady( or (Ophelia(1 Wild roses have a delicate
aroma0 unmistakale0 ut not to:s%oon:for like ("emorial 'ay( or (EliFaeth )arkness( or (,ra!rant
Cloud(1 The human eye and the human nose %ent to %ork on %ild roses0 enlar!in! them0 shapin!
them0 doulin! up the petals0 tintin! them0 refinin! the loom0 oostin! natural fra!rances to heady
extremes0 adHustin! haits of !ro%th0 eventually enterin! them in sophisticated hyridiFation
pro!rams until0 today0 after decades of skilful selective reedin!0 there are hundreds of priFed
varieties0 each %ith its o%n evocative or commemorative name1 Who %ould not like to have a rose
named after herB
Roses tell the same story as do!s0 ut %ith one difference0 %hich is relevant to our
softenin!:up strate!y1 The flo%er of the rose0 even efore human eyes and noses emarked on their
%ork of !enetic chisellin!0 o%ed its very existence to millions of years of very similar sculptin! y
insect eyes and noses 7%ell0 antennae0 %hich is %hat insects smell %ith81 &nd the same is true of all
the flo%ers that eautify our !ardens1
The sunflo%er0 )elianth"s ann""s0 is a *orth &merican plant %hose %ild form looks like an
aster or lar!e daisy1 Cultivated sunflo%ers today have een domesticated to the point %here their
flo%ers are the siFe of a dinner plate1 L ("ammoth( sunflo%ers0 ori!inally red in Russia0 are 23 to
2@ feet hi!h0 the head diameter is close to one foot0 %hich is more than ten times the siFe of a %ild
sunflo%er(s disc0 and there is normally only one head per plant0 instead of the many0 much smaller0
flo%ers of the %ild plant1 The Russians started reedin! this &merican flo%er0 y the %ay0 for
reli!ious reasons1 'urin! -ent and &dvent0 the use of oil in cookin! %as anned y the Orthodox
Church1 Conveniently0 and for a reason that # : untutored in the profundities of theolo!y : shall not
presume to fathom0 sunflo%er seed oil %as deemed to e exempt from this prohiition1 O This
provided one of the economic pressures that drove the recent selective reedin! of the sunflo%er1
-on! efore the modern era0 ho%ever0 native &mericans had een cultivatin! these nutritious and
spectacular flo%ers for food0 for dyes and for decoration0 and they achieved results intermediate
et%een the %ild sunflo%er and the extrava!ant extremes of modern cultivars1 But efore that
a!ain0 sunflo%ers0 like all ri!htly coloured flo%ers0 o%ed their very existence to selective reedin!
y insects1
The same is true of most of the flo%ers %e are a%are of : proaly all the flo%ers that are
coloured anythin! other than !reen and %hose smell is anythin! more than Hust va!uely plant:like1
*ot all the %ork %as done y insects : for some flo%ers the pollinators that did the initial selective
reedin! %ere hummin!irds0 ats0 even fro!s : ut the principle is the same1 Garden flo%ers have
een further enhanced y us0 ut the %ild flo%ers %ith %hich %e started only cau!ht our attention
in the first place ecause insects and other selective a!ents had een there efore us1 Generations of
ancestral flo%ers %ere chosen y !enerations of ancestral insects or hummin!irds or other natural
pollinators1 #t is a perfectly !ood example of selective reedin!0 %ith the minor difference that the
reeders %ere insects and hummin!irds0 not humans1 &t least0 # think the difference is minor1 6ou
may not0 in %hich case # still have some softenin! up to do1
What mi!ht tempt us to think it a maHor differenceB ,or one thin!0 humans conscio"sly set
out to reed0 say0 the darkest0 most lackish purple rose they can0 and they do it to satisfy an
aesthetic %him0 or ecause they think other people %ill pay money for it1 #nsects do it not for
aesthetic reasons ut for reasons of 1 1 1 %ell0 here %e need to ack up and look at the %hole matter
of flo%ers and their relationship %ith their pollinators1 )ere(s the ack!round1 ,or reasons # %on(t
!o into no%0 it is of the essence of sexual reproduction that you shouldn(t fertiliFe yourself1 #f you
did that0 after all0 there(d e little point in otherin! %ith sexual reproduction in the first place1
Pollen must someho% e transported from one plant to another1 )ermaphroditic plants that have
male and female parts %ithin one flo%er often !o to elaorate len!ths to stop the male half from
fertiliFin! the female half1 'ar%in himself studied the in!enious %ay this is achieved in primroses1
Takin! the need for cross:fertiliFation as a !iven0 ho% do flo%ers achieve the feat of movin!
pollen across the physical !ap that separates them from other flo%ers of the same speciesB The
ovious %ay is y the %ind0 and plenty of plants use it1 Pollen is a fine0 li!ht po%der1 #f you release
enou!h of it on a reeFy day0 one or t%o !rains may have the luck to land on the ri!ht spot in a
flo%er of the ri!ht species1 But %ind pollination is %asteful1 & hu!e surplus of pollen needs to e
manufactured0 as hay fever sufferers kno%1 The vast maHority of pollen !rains land some%here
other than %here they should0 and all that ener!y and costly materiel is %asted1 There is a more
directed %ay for pollen to e tar!eted1
Why don(t plants choose the animal option0 and %alk around lookin! for another plant of the
same species0 then copulate %ith itB That(s a harder Euestion to deal %ith than you mi!ht think1 #t(s
circular simply to assert that plants don(t %alk0 ut #(m afraid that %ill have to do for no%1L The fact
is0 plants don(t %alk1 But animals %alk1 &nd animals fly0 and they have nervous systems capale of
directin! them to%ards particular tar!ets0 %ith sou!ht:for shapes and colours1 So if only there %ere
some %ay to persuade an animal to dust itself %ith pollen and then %alk or preferaly fly to another
plant of the ri!ht species 1 1 1
Well0 the ans%er(s no secretA that(s exactly %hat happens1 The story is in some cases hi!hly
complex and in all cases fascinatin!1 "any flo%ers use a rie of food0 usually nectar1 "aye rie
is too loaded a %ord1 Would you prefer (payment for services rendered(B #(m happy %ith oth0 so
lon! as %e don(t misunderstand them in a human %ay1 *ectar is su!ary syrup0 and it is
manufactured y plants specifically and only for payin!0 and fuellin!0 ees0 utterflies0
hummin!irds0 ats and other hired transport1 #t is costly to make0 funnellin! off a proportion of the
sunshine ener!y trapped y the leaves0 the solar panels of the plant1 ,rom the point of vie% of the
ees and hummin!irds0 it is hi!h:ener!y aviation fuel1 The ener!y locked up in the su!ars of nectar
could have een used else%here in the economy of the plant0 perhaps to make roots0 or to fill the
under!round stora!e ma!aFines that %e call tuers0 uls and corms0 or even to make hu!e
Euantities of pollen for roadcastin! to the four %inds1 Evidently0 for a lar!e numer of plant
species0 the trade:off %orks out in favour of payin! insects and irds for their %in!s0 and fuellin!
their fli!ht muscles %ith su!ar1 #t(s not a totally over%helmin! advanta!e0 ho%ever0 ecause some
plants do use %ind pollination0 presumaly ecause details of their economic circumstances tip their
alance that %ay1 Plants have an ener!y economy and0 as %ith any economy0 trade:offs may favour
different options under different circumstances1 That(s an important lesson in evolution0 y the %ay1
'ifferent species do thin!s in different %ays0 and %e often %on(t understand the differences until
%e have examined the %hole economy of the species1
#f %ind pollination is at one end of a continuum of cross:fertiliFation techniEues : shall %e
call it the profli!ate endB : %hat is at the other end0 the (ma!ic ullet( endB +ery fe% insects can e
relied upon to fly like a ma!ic ullet strai!ht from the flo%er %here they have picked up pollen to
another flo%er of exactly the ri!ht species1 Some Hust !o to any old flo%er0 or possily any flo%er
of the ri!ht colour0 and it is still a matter of luck %hether it happens to e the same species as the
flo%er that has Hust paid it in nectar1 *evertheless0 there are some lovely examples of flo%ers that
lie far out to%ards the ma!ic ullet end of the continuum1 )i!h on the list are orchids0 and it(s no
%onder that 'ar%in devoted a %hole ook to them1
Both 'ar%in and his co:discoverer of natural selection0 Wallace0 called attention to an
amaFin! orchid from "ada!ascar0 An$raec"m ses3"ipedale 7see colour pa!e >80 and oth men
made the same remarkale prediction0 %hich %as later triumphantly vindicated1 This orchid has
tuular nectaries that reach do%n more than 22 inches y 'ar%in(s o%n ruler1 That(s nearly 45
centimetres1 & related species0 An$raec"m lon$icalcar0 has nectar:earin! spurs that are even
lon!er0 up to >5 centimetres 7more than 2= inches81 'ar%in0 purely on the stren!th of A4
ses3"ipedale(s existence in "ada!ascar0 predicted in his orchid ook of 2;<3 that there must e
(moths capale of extension to a len!th of et%een ten and eleven inches(1 Wallace0 five years later
7it isn(t clear %hether he had read 'ar%in(s ook8 mentioned several moths %hose proosces %ere
nearly lon! enou!h to meet the case1
# have carefully measured the prooscis of a specimen of %acrosila cl"enti"s from South
&merica in the collection of the British "useum0 and find it to e nine inches and a Euarter lon!K
One from tropical &frica 7 %acrosila mor$anii8 is seven inches and a half1 & species havin! a
prooscis t%o or three inches lon!er could reach the nectar in the lar!est flo%ers of An$raec"m
ses3"ipedale0 %hose nectaries vary in len!th from ten to fourteen inches1 That such a moth exists in
"ada!ascar may e safely predictedJ and naturalists %ho visit that island should search for it %ith
as much confidence as astronomers searched for the planet *eptune0 : and they %ill e eEually
#n 29540 after 'ar%in(s death ut %ell %ithin Wallace(s lon! lifetime0 a hitherto unkno%n
moth %as discovered %hich turned out to fulfil the 'ar%inGWallace prediction0 and %as duly
honoured %ith the su:specific name praedicta1 But even 5anthopan mor$ani praedicta0 ('ar%in(s
ha%k moth(0 is not sufficiently %ell endo%ed to pollinate A4 lon$icalcar0 and the existence of this
flo%er encoura!es us to suspect the existence of an even lon!er:ton!ued moth0 %ith the same
confidence as Wallace invoked the predicted discovery of the planet *eptune1 By the %ay0 this little
example !ives the lie0 yet a!ain0 to the alle!ation that evolutionary science cannot e predictive
ecause it concerns past history1 The 'ar%inGWallace prediction %as still a perfectly valid one0
even thou!h the praedicta moth must already have existed efore they made it1 They %ere
predictin! that0 at some time in the future0 someody %ould discover a moth %ith a ton!ue lon!
enou!h to reach the nectar in A4 ses3"ipedale1
#nsects have !ood colour vision0 ut their %hole spectrum is shifted to%ards the ultraviolet
and a%ay from the red1 -ike us0 they see yello%0 !reen0 lue and violet1 $nlike us0 ho%ever0 they
also see %ell into the ultraviolet ran!eJ and they don(t see red0 at (our( end of the spectrum1 #f you
have a red tuular flo%er in your !arden it is a !ood et0 thou!h not a certain prediction0 that in the
%ild it is pollinated not y insects ut y irds0 %ho see %ell at the red end of the spectrum :
perhaps hummin!irds if it is a *e% World plant0 or sunirds if an Old World plant1 ,lo%ers that
look plain to us may actually e lavishly decorated %ith spots or stripes for the enefit of insects0
ornamentation that %e can(t see ecause %e are lind to ultraviolet1 "any flo%ers !uide ees in to
land y little run%ay markin!s0 painted on the flo%er in ultraviolet pi!ments0 %hich the human eye
can(t see1
The evenin! primrose 7!enothera8 looks yello% to us1 But a photo!raph taken throu!h an
ultraviolet filter sho%s a pattern for the enefit of ees0 %hich %e can(t see %ith normal vision 7see
colour pa!e =81 #n the photo!raph it appears as red0 ut that is a (false colour(A an aritrary choice y
the photo!raphic process1 #t doesn(t mean that ees %ould see it as red1 *oody kno%s ho%
ultraviolet 7or yello% or any other colour8 looks to a ee 7# don(t even kno% ho% red looks to you :
an old philosophical chestnut81
& meado% full of flo%ers is nature(s Times SEuare0 nature(s Piccadilly Circus1 & slo%:
motion neon si!n0 it chan!es from %eek to %eek as different flo%ers come into season0 carefully
prompted y cues from0 for example0 the chan!in! len!th of days to synchroniFe %ith others of
their o%n species1 This floral extrava!anFa0 splashed across the !reen canvas of a meado%0 has een
shaped and coloured0 ma!nified and titivated y the past choices made y animal eyesA ee eyes0
utterfly eyes0 hoverfly eyes1 #n *e% World forests %e(d have to add hummin!ird or in &frican
forests sunird eyes to the list1
)ummin!irds and sunirds are not particularly closely related0 y the %ay1 They look and
ehave like each other ecause they have conver!ed upon the same %ay of life0 lar!ely revolvin!
around flo%ers and nectar 7althou!h they eat insects as %ell as nectar81 They have lon! eaks for
proin! nectaries0 extended y even lon!er ton!ues1 Sunirds are less accomplished hoverers than
hummin!irds0 %ho can even !o ack%ards like a helicopter1 &lso conver!ent0 althou!h from a far
distant vanta!e point in the animal kin!dom0 are the hummin!ird ha%k moths0 a!ain consummate
hoverers %ith spectacularly lon! ton!ues 7all three types of nectar Hunkie are illustrated on colour
pa!e =81
We shall return to conver!ent evolution later in the ook0 after properly understandin!
natural selection1 )ere0 in this chapter0 flo%ers are seducin! us0 dra%in! us in0 step y step0 linin!
our path to that understandin!1 )ummin!ird eyes0 ha%k:moth eyes0 utterfly eyes0 hoverfly eyes0
ee eyes are critically cast over %ild flo%ers0 !eneration after !eneration0 shapin! them0 colourin!
them0 s%ellin! them0 patternin! and stipplin! them0 in almost exactly the same %ay as human eyes
later did %ith our !arden varietiesJ and %ith do!s0 co%s0 caa!es and corn1
,or the flo%er0 insect pollination represents a hu!e advance in economy over the %asteful
scatter!un of %ind pollination1 Even if a ee visits flo%ers indiscriminately0 lurchin! promiscuously
from uttercup to cornflo%er0 from poppy to celandine0 a pollen !rain clin!in! to its hairy adomen
has a much !reater chance of hittin! the ri!ht tar!et : a second flo%er of the same species : than it
%ould have if scattered on the %ind1 Sli!htly etter %ould e a ee %ith a preference for a particular
colour0 say lue1 Or a ee that0 %hile not havin! any lon!:term colour preference0 tends to form
colour haits0 so that it chooses colours in runs1 Better still %ould e an insect that visits flo%ers of
only one species1 &nd there are flo%ers0 like the "ada!ascar orchid that inspired the
'ar%inGWallace prediction0 %hose nectar is availale only to certain insects that specialiFe in that
kind of flo%er and enefit from their monopoly over it1 Those "ada!ascar moths are the ultimate
ma!ic ullets1
,rom a moth(s point of vie%0 flo%ers that relialy provide nectar are like docile0 productive
milch co%s1 ,rom the flo%ers( point of vie%0 moths that relialy transport their pollen to other
flo%ers of the same species are like a %ell:paid ,ederal Express service0 or like %ell:trained homin!
pi!eons1 Each side could e said to have domesticated the other0 selectively reedin! them to do a
etter Ho than they previously did1 )uman reeders of priFe roses have had almost exactly the same
kinds of effects on flo%ers as insects have : Hust exa!!erated them a it1 #nsects red flo%ers to e
ri!ht and sho%y1 Gardeners made them ri!hter and sho%ier still1 #nsects made roses pleasantly
fra!rant1 We came alon! and made them even more so1 #ncidentally0 it is a fortunate coincidence
that the fra!rances that ees and utterflies prefer happen to appeal to us too1 ,lo%ers such as
(stinkin! BenHamin( 7 Trilli"m erect"m8 or the (corpse flo%er( 7 Amorphophall"s titan"m80 %hich use
flesh flies or carrion eetles as pollinators0 often nauseate us0 ecause they mimic the smell of
decayin! meat1 Such flo%ers have not0 # presume0 had their scents enhanced y human
Of course0 the relationship et%een insects and flo%ers is a t%o:%ay street0 and %e mustn(t
ne!lect to look in oth directions1 #nsects may (reed( flo%ers to e more eautiful0 ut not ecause
they enHoy the eauty1L Rather0 the flo%ers enefit from ein! perceived as attractive y insects1
The insects0 y choosin! the most attractive flo%ers to visit0 inadvertently (reed for( floral eauty1
&t the same time0 the flo%ers are reedin! the insects for pollination aility1 Then a!ain0 # have
implied that insects reed flo%ers for hi!h nectar yield0 like dairymen reedin! massively uddered
,riesians1 But it is in the flo%ers( interests to ration their nectar1 Satiate an insect and it has no
incentive to !o on and look for a second flo%er : ad ne%s for the first flo%er0 for %hich the second
visit0 the pollinatin! visit0 is the %hole point of the exercise1 ,rom the flo%ers( point of vie%0 a
delicate alance must e struck et%een providin! too much nectar 7no visit to a second flo%er8 and
too little 7no incentive to visit the first flo%er81
#nsects have milked flo%ers for their nectar0 and red them for increased yield : proaly
encounterin! resistance from the flo%ers0 as %e have Hust seen1 )ave eekeepers 7or
horticulturalists %ith the interests of eekeepers in mind8 red flo%ers to e even more productive
of nectar0 Hust as dairy farmers red ,riesian and .ersey co%sB #(d e intri!ued to kno% the ans%er1
"ean%hile0 there(s no dout of the close parallel et%een horticulturalists as reeders of pretty and
fra!rant flo%ers0 and ees and utterflies0 hummin!irds and sunirds doin! the same thin!1
6O$ &RE "6 *&T$R&- SE-ECT#O*
&re there other examples of selective reedin! y non:human eyesB Oh yes1 Think of the
dull0 camoufla!ed pluma!e of a hen pheasant0 compared %ith the splendiferous male of the same
species1 There seems little dout that0 if his individual survival %ere the only thin! that mattered0
the cock !olden pheasant %ould (prefer( to look like the female0 or like a !ro%n:up version of ho%
he %as as a chick1 The female and the chicks are oviously %ell camoufla!ed0 and that(s the %ay the
male %ould e too if individual survival %ere his priority1 The same is true of other pheasants such
as -ady &mherst(s and the familiar rin!:necked pheasant1 The cocks look flamoyant and
dan!erously attractive to predators0 ut each species in a very different %ay1 The hens are
camoufla!ed and dull:coloured0 each species in pretty much the same %ay1 What is !oin! on hereB
One %ay to put it is 'ar%in(s %ayA (sexual selection(1 But another %ay : and the one that
etter suits my primrose path : is (selective reedin! y females of males(1 Bri!ht colours may
indeed attract predators0 ut they attract female pheasants too1 Generations of hens chose to mate
%ith ri!ht0 !lo%in! males0 rather than the dull ro%n creatures that the males %ould surely have
remained ut for selective reedin! y females1 The same thin! happened %ith peahens selectively
reedin! peacocks0 female irds of paradise reedin! males0 and numerous other examples of irds0
mammals0 fish0 amphiians0 reptiles and insects %here females 7it(s usually females rather than
males0 for reasons %e needn(t !o into8 choose from amon! competin! males1 &s %ith !arden
flo%ers0 human pheasant:reeders have improved upon the selective handi%ork of the hen
pheasants that preceded them0 producin! spectacular variants of the !olden pheasant0 for example0
althou!h more y pickin! one or t%o maHor mutations rather than y !radually shapin! the ird
throu!h !enerations of reedin!1 )umans have also selectively red some amaFin! varieties of
pi!eons 7as 'ar%in kne% at first hand8 and chickens0 descended from a ,ar Eastern ird0 the red
Hun!le fo%l Gall"s $all"s4
+arieties o$ hi&en: three ill(strations $ro" Dar,in's The Variation of Animals and
Plants under Domestication
This chapter is mostly aout selection y eyes0 ut other senses can do the same thin!1
,anciers have red canaries for their son!s0 as %ell as for their appearance1 The %ild canary is a
yello%ish ro%n finch0 not spectacular to look at1 )uman selective reeders have taken the palette
of colours thro%n up y random !enetic variation and manufactured a colour distinctive enou!h to
e named after the irdA canary yello%1 By the %ay0 the ird itself is named after the islands0L not
the other %ay around as %ith the Galapa!os #slands0 %hose name comes from a Spanish %ord for
tortoise1 But canaries are est kno%n for their son!0 and this too has een tuned up and enriched y
human reeders1 +arious son!sters have een manufactured0 includin! Rollers0 %hich have een
red to sin! %ith the eak closed0 Watersla!ers0 %hich sound like ulin! %ater0 and Timrados0
%hich produce metallic0 ell:like notes0 to!ether %ith a castanet:like chatter that efits their Spanish
ori!ins1 'omestically red son!s are lon!er0 louder and more freEuent than the %ild ancestral type1
But all these hi!hly priFed son!s are made up of elements that occur in %ild canaries0 Hust as the
haits and tricks of various reeds of do!s come from elements to e found in the ehavioural
repertoire of %olves1 L
Once a!ain0 human reeders have only een uildin! on the earlier selective reedin! efforts
of female irds1 Over !enerations0 %ild female canaries inadvertently red males for their sin!in!
pro%ess y choosin! to mate %ith males %hose son!s %ere especially appealin!1 #n the particular
case of canaries it happens that %e kno% a little more1 Canaries 7and Barary doves8 have een
favourite suHects for research on hormones and reproductive ehaviour1 #t is kno%n that in oth
species the sound of male vocaliFation 7even from a tape recordin!8 causes the females( ovaries to
s%ell and secrete hormones that rin! them into reproductive condition and make them more ready
to mate1 One could say that male canaries are manipulatin! females y sin!in! to them1 #t is almost
as thou!h they %ere !ivin! them hormone inHections1 One could also say that females are
selectively reedin! males to ecome etter and etter at sin!in!1 The t%o %ays of lookin! at the
matter are t%o sides of the same coin1 &s %ith other ird species0 y the %ay0 there is a
complicationA son! is not only appealin! to females0 it is also a deterrent to rival males : ut #(ll
leave that on one side1
*o%0 to move the ar!ument on0 look at the pictures opposite1 The first is a %oodcut of a
.apanese kauki mask0 representin! a samurai %arrior1 The second is a cra of the species )eikea
6aponica0 %hich is found in .apanese %aters1 The !eneric name0 )eikea0 comes from a .apanese
clan called the )eike0 %ho %ere defeated at sea in the attle of 'anno:$ra 722;=8 y a rival clan
called the GenHi1 -e!end tells that the !hosts of dro%ned )eike %arriors no% inhait the ottom of
the sea0 in the odies of cras : )eikea 6aponica1 The myth is encoura!ed y the pattern on the ack
of this cra0 %hich resemles the fiercely !rimacin! face of a samurai %arrior1 The famous
Foolo!ist Sir .ulian )uxley %as impressed enou!h y the resemlance to %rite0 (The resemlance of
Dorippe to an an!ry .apanese %arrior is far too specific and far too detailed to e accidental 1 1 1 #t
came aout ecause those cras %ith a more perfect resemlance to a %arrior(s face %ere less
freEuently eaten than the others1( 7 Dorippe %as %hat the cra %as called in 29=3 %hen )uxley
%rote1 #t reverted to )eikea in 2995 %hen someody rediscovered that it had een so named as
early as 2;3> : such are the strict priority rules of Foolo!ical nomenclature18
Ka*(&i "as& o$ sa"(rai ,arrior
Heikea japonica ra*
This theory0 that !enerations of superstitious fishermen thre% ack into the sea cras that
resemled human faces0 received ne% le!s in 29;5 %hen Carl Sa!an discussed it in his %onderful
#osmos1 #n his %ords0
Suppose that0 y chance0 amon! the distant ancestors of this cra0 one arose that resemled0
even sli!htly0 a human face1 Even efore the attle of 'anno:ura0 fishermen may have een
reluctant to eat such a cra1 #n thro%in! it ack0 they set in motion an evolutionary process 1 1 1 &s
the !enerations passed0 of cras and fishermen alike0 the cras %ith patterns that most resemled a
samurai face survived preferentially until eventually there %as produced not Hust a human face0 not
Hust a .apanese face0 ut the visa!e of a fierce and sco%lin! samurai1
#t(s a lovely theory0 too !ood to die easily0 and the meme has indeed replicated itself throu!h
the canon1 # even found a %esite %here you can vote on %hether the theory is true 742 per cent of
20442 voters80 %hether the photo!raphs are fakes 72= per cent80 %hether .apanese craftsmen carve
the shells to look that %ay 7< per cent80 %hether the resemlance is Hust a coincidence 74; per cent80
or even %hether the cras really are manifestations of dro%ned samurai %arriors 7an amaFin! 25 per
cent81 Scientific truths are not0 of course0 decided y pleiscite0 and # voted only ecause # %as
other%ise not allo%ed to see the votin! fi!ures1 #(m afraid # voted %ith the killHoys1 # think0 on
alance0 that the resemlance is proaly a coincidence1 *ot ecause0 as one authoritative sceptic
has pointed out0 the rid!es and !rooves on the cra(s ack actually si!nify underlyin! muscle
attachments1 Even on the )uxleyGSa!an theory0 the superstitious fishermen %ould have to have
e!un y noticin! some kind of ori!inal resemlance0 ho%ever sli!ht0 and a symmetrical pattern of
muscle attachments is exactly the kind of thin! that %ould have provided that initial resemlance1 #
am more impressed y the same sceptic(s oservation that these cras are too small to e %orth
eatin! any%ay1 &ccordin! to him0 all cras of that siFe %ould have een thro%n ack0 %hether or
not their acks resemled human faces0 althou!h # have to say that this more tellin! source of
scepticism had a lar!e ite taken out of it %hen # %as taken out to dinner in Tokyo and my host
ordered0 for all the company0 a dish of cras1 They %ere much lar!er than )eikea0 and they %ere
thickly encrusted in stout0 calcified carapaces0 ut that didn(t stop this superman pickin! up %hole
cras0 one y one0 and itin! into them like an apple0 %ith a crunchin! sound that seemed to presa!e
hideously leedin! !ums1 & cra as small as )eikea %ould e a doddle to such a !astronomic
champion1 )e %ould surely s%allo% it %hole %ithout attin! an eyelid1
"y main reason for scepticism aout the )uxleyGSa!an theory is that the human rain is
demonstraly ea!er to see faces in random patterns0 as %e kno% from scientific evidence0 on top of
the numerous le!ends aout faces of .esus0 or the +ir!in "ary0 or "other Teresa0 ein! seen on
slices of toast0 or piFFas0 or patches of damp on a %all1 This ea!erness is enhanced if the pattern
departs from randomness in the specific direction of ein! symmetrical1 &ll cras 7except hermit
cras8 are symmetrical any%ay1 # reluctantly suspect that the resemlance of )eikea to a samurai
%arrior is no more than an accident0 much as # %ould like to elieve it has een enhanced y natural
*ever mind1 There are plenty of other examples not involvin! humans0 %here animal
(fishermen(0 as it %ere0 (thro% ack( 7or don(t see in the first place8 %ould:e food ecause of a
resemlance to somethin! sinister0 and %here the resemlance is certainly not due to chance1 #f you
%ere a ird0 out huntin! caterpillars in the forest0 %hat %ould you do if you %ere suddenly
confronted %ith a snakeB -eap ack startled0 %ould e my !uess0 and then !ive it a %ide erth1
Well0 there is a caterpillar : to e precise0 the rear end of a caterpillar : that ears an unmistakale
resemlance to a snake1 #t is truly alarmin! if you are fri!htened of snakes : as # shamefacedly
confess # am1 # even think # mi!ht e reluctant to pick this animal up0 despite kno%in! perfectly
%ell that it is in fact a harmless caterpillar1 7& picture of this extraordinary creature appears on
colour pa!e @18 # have the same prolem %ith pickin! up %asp:mimickin! or ee:mimickin!
hoverflies0 even thou!h # can see0 from their possession of only one pair of %in!s0 that they are
stin!less flies1 These are amon! a vast list of animals that !ain protection ecause they look like
somethin! elseA somethin! inedile like a pele0 a t%i! or a frond of sea%eed0 or somethin!
positively nasty like a snake or a %asp or the !larin! eyes of a possile predator1
)ave ird eyes0 then0 een reedin! insects for their resemlance to unpalatale or
venomous modelsB There(s one sense in %hich %e surely have to ans%er yes1 What0 after all0 is the
difference et%een this and peahens reedin! peacocks for eauty0 or humans reedin! do!s or
rosesB "ainly0 peahens are reedin! positiely for somethin! attractive0 y approachin! it0 %hile the
caterpillar:huntin! irds are reedin! ne$atiely for somethin! repellent0 y avoidin! it1 Ri!ht then0
here(s another example0 and in this case the (reedin!( is positive0 even thou!h the selector doesn(t
enefit from its choice1 ,ar from it1
'eep:sea an!ler fish sit on the ottom of the sea0 %aitin! patiently for prey1L -ike many
deep:sea fish0 an!lers are spectacularly u!ly y our standards1 "aye y fish standards too0
althou!h it proaly doesn(t matter ecause0 do%n %here they live0 it is too dark to see much
any%ay1 -ike other deniFens of the deep sea0 female an!ler fish often make their o%n li!ht : or
rather0 they have special receptacles in %hich they house acteria %hich make li!ht for them1 Such
(ioluminescence( isn(t ri!ht enou!h to reflect any detail0 ut it is ri!ht enou!h to attract other
fish1 & spine %hich0 in a normal fish0 %ould e Hust one of the rays in a fin0 ecomes elon!ated and
stiffened to make a fishin! rod1 #n some species the (rod( is so lon! and flexile that you(d call it a
line rather than a rod1 &nd on the end of the fishin! rod or line is : %hat elseB : a ait0 or lure1 The
aits vary from species to species0 ut they al%ays resemle small food itemsA perhaps a %orm0 or a
small fish0 or Hust a nondescript ut temptin!ly Hi!!lin! morsel1 Often the ait is actually luminousA
another natural neon si!n0 and in this case the messa!e ein! flashed is (come and eat me(1 Small
fish are indeed tempted1 They approach close to the ait1 &nd it is the last thin! they do for0 at that
moment0 the an!ler opens her hu!e ma% and the prey is en!ulfed %ith the inrush of %ater1
*o%0 %ould %e say that the small prey fish are (reedin! for( more and more appealin!
lures0 Hust as peahens reed for more appealin! peacocks0 and horticulturalists reed for more
appealin! rosesB #t(s hard to see %hy not1 #n the case of the roses0 the most attractive looms are the
ones delierately chosen for reedin! y the !ardener1 "uch the same is true of peacocks chosen y
peahens1 #t is possile that the peahens are not a%are that they are choosin!0 %hereas the rose:
!ro%ers are1 But that doesn(t seem a very important distinction under the circumstances1 Sli!htly
more compellin! is a distinction et%een the an!ler fish example and the other t%o1 The prey fish
are indeed choosin! the most (attractive( an!ler fish for reedin!0 via the indirect route of choosin!
them for survival y feedin! themK &n!lers %ith unattractive lures are more likely to starve to death
and therefore less likely to reed1 &nd the small prey fish are indeed doin! the (choosin!(1 But they
are choosin! %ith their livesK What %e are homin! in on here is true natural selection0 and %e are
reachin! the end of the pro!ressive seduction that is this chapter1
)ere(s the pro!ression laid out1
2 )umans delierately choose attractive roses0 sunflo%ers etc1 for reedin!0 therey
preservin! the !enes that produce the attractive features1 This is called artificial selection0 it(s
somethin! humans have kno%n aout since lon! efore 'ar%in0 and everyody understands that it
is po%erful enou!h to turn %olves into chihuahuas and to stretch maiFe cos from inches to feet1
3 Peahens 7%e don(t kno% %hether consciously and delierately0 ut let(s !uess not8 choose
attractive peacocks for reedin!0 a!ain therey preservin! attractive !enes1 This is called sexual
selection0 and 'ar%in discovered it0 or at least clearly reco!niFed it and named it1
4 Small prey fish 7definitely not delierately8 choose attractive an!ler fish for survival0 y
feedin! the most attractive ones %ith their o%n odies0 therey inadvertently choosin! them for
reedin! and passin! on0 and therefore preservin!0 the !enes that produce the attractive features1
This is called : yes0 %e(ve finally !ot there : natural selection0 and it %as 'ar%in(s !reatest
'ar%in(s special !enius realiFed that nature could play the role of selectin! a!ent1
Everyody kne% aout artificial selection0L or at least everyody %ith any experience of farms or
!ardens0 do! sho%s or dovecotes1 But it %as 'ar%in %ho first spotted that you don(t have to have a
choosin! a$ent1 The choice can e made automatically y survival : or failure to survive1 Survival
counts0 'ar%in realiFed0 ecause only survivors reproduce and pass on the !enes 7'ar%in didn(t use
the %ord8 that helped them to survive1
# chose the an!ler fish as my example0 ecause this can still e represented as an a!ent usin!
its eyes to choose that %hich survives1 But %e have reached the point in our ar!ument : 'ar%in(s
point : %here %e no lon!er need to talk aout a choosin! a!ent at all1 "ove no% from an!ler fish
to0 say0 tuna or tarpon0 fish that actively pursue their prey1 By no sensile stretch of lan!ua!e or
ima!ination could %e claim that the prey (choose( %hich tarpon survive y ein! eaten1 What %e
can say0 ho%ever0 is that the tarpon that are etter eEuipped to catch prey0 for %hatever reason : fast
s%immin! muscles0 keen eyes0 etc1 : %ill e the ones that survive0 and therefore the ones that
reproduce and pass on the !enes that made them successful1 They are (chosen( y the very act of
stayin! alive0 %hereas another tarpon that %as0 for whateer reason0 less %ell eEuipped %ould not
survive1 So0 %e can add a fourth step to our list1
> Without any kind of choosin! a!ent0 those individuals that are (chosen( y the fact that
they happen to possess superior eEuipment to survive are the most likely to reproduce0 and therefore
to pass on the !enes for possessin! superior eEuipment1 Therefore every !ene pool0 in every species0
tends to ecome filled %ith !enes for makin! superior eEuipment for survival and reproduction1
*otice ho% all:encompassin! natural selection is1 The other examples # have mentioned0
steps 20 3 and 4 and lots of others0 can all e %rapped up in natural selection0 as special cases of the
more !eneral phenomenon1 'ar%in %orked out the most !eneral case of a phenomenon that people
already kne% aout in restricted form1 )itherto0 they had kno%n aout it only in the special case of
artificial selection1 The !eneral case is the non:random survival of randomly varyin! hereditary
eEuipment1 #t doesn(t matter ho% the non:random survival comes aout1 #t can e delierate0
explicitly intentional choice y an a!ent 7as %ith humans choosin! pedi!ree !reyhounds for
reedin!8J it can e inadvertent choice y an a!ent %ithout explicit intention 7as %ith peahens
choosin! peacocks for reedin!8J it can e inadvertent choice %hich the chooser : %ith a hindsi!ht
that is !ranted to us ut not the chooser itself : %ould prefer not to have made 7as %ith prey fish
choosin! to approach an an!ler fish(s lure8J or it can e somethin! that %e %ouldn(t reco!niFe as
choice at all0 as %hen a tarpon survives y virtue of0 say0 an oscure iochemical advanta!e uried
deep %ithin its muscles0 %hich !ives it an extra urst of speed %hen pursuin! prey1 'ar%in himself
said it eautifully0 in a favourite passa!e from !n the !ri$in of SpeciesA
#t may e said that natural selection is daily and hourly scrutinisin!0 throu!hout the %orld0
every variation0 even the sli!htestJ reHectin! that %hich is ad0 preservin! and addin! up all that is
!oodJ silently and insensily %orkin!0 %henever and %herever opportunity offers0 at the
improvement of each or!anic ein! in relation to its or!anic and inor!anic conditions of life1 We
see nothin! of these slo% chan!es in pro!ress0 until the hand of time has marked the lon! lapse of
a!es0 and then so imperfect is our vie% into lon! past !eolo!ical a!es0 that %e see only that the
forms of life are no% different from %hat they formerly %ere1
# have here Euoted0 as is my usual practice0 the first edition of 'ar%in(s masterpiece1 &n
interestin! interpolation found its %ay into later editionsA (#t may metaphorically e said that natural
selection is daily and hourly 1 1 1( 7emphasis added81 6ou mi!ht think that (#t may e said 1 1 1( %as
cautious enou!h1 But in 2;<< 'ar%in received a letter from Wallace0 co:discoverer of natural
selection0 su!!estin! that an even hi!her hed!e a!ainst misunderstandin! %as re!rettaly necessary1
"y dear 'ar%in0 : # have een so repeatedly struck y the utter inaility of numers of
intelli!ent persons to see clearly0 or at all0 the self:actin! and necessary effects of *atural Selection0
that # am led to conclude that the term itself0 and your mode of illustratin! it0 ho%ever clear and
eautiful to many of us0 are yet not the est adapted to impress it on the !eneral naturalist pulic1
Wallace %ent on to Euote a ,rench author called .anet0 %ho %as evidently0 unlike Wallace
and 'ar%in0 a deeply muddled individualA
# see that he considers your %eak point to e that you do not see that (thou!ht and direction
are essential to the action of *atural Selection1( The same oHection has een made a score of times
y your chief opponents0 and # have heard it as often stated myself in conversation1 *o%0 # think
this arises almost entirely from your choice of the term *atural Selection0 and so constantly
comparin! it in its effects to man(s selection0 and also to your so freEuently personifyin! nature as
(selectin!(0 as (preferrin!( 1 1 1 etc10 etc1 To the fe% this is as clear as dayli!ht0 and eautifully
su!!estive0 ut to many it is evidently a stumlin!:lock1 # %ish0 therefore0 to su!!est to you the
possiility of entirely avoidin! this source of misconception in your !reat %ork0 and also in future
editions of the (Ori!in0( and # think it may e done %ithout difficulty and very effectually y
adoptin! Spencer(s term 1 1 1 (Survival of the ,ittest1( This term is the plain expression of the factJ
(*atural Selection( is a metaphorical expression of it 1 1 1
Wallace had a point1 $nfortunately0 Spencer(s term (Survival of the ,ittest( raises prolems
of its o%n0 %hich Wallace couldn(t have foreseen0 and # %on(t !o into them here1 #n spite of
Wallace(s %arnin!0 # prefer to follo% 'ar%in(s o%n strate!y of introducin! natural selection via
domestication and artificial selection1 # like to think that "onsieur .anet mi!ht have !ot the point
this time around1 But # did have another reason0 too0 for follo%in! 'ar%in(s lead0 and it is a !ood
one1 The ultimate test of a scientific hypothesis is experiment1 Experiment specifically means that
you don(t Hust %ait for nature to do somethin!0 and passively oserve it and see %hat it correlates
%ith1 6ou !o in there and do somethin!1 6ou manip"late1 6ou chan$e somethin!0 in a systematic
%ay0 and compare the result %ith a (control( that lacks the chan!e0 or you compare it %ith a different
Experimental interference is of enormous importance0 ecause %ithout it you can never e
sure that a correlation you oserve has any causal si!nificance1 This can e illustrated y the so:
called (church clocks fallacy(1 The clocks in the to%ers of t%o nei!hourin! churches chime the
hours0 ut St &(s a little efore St B(s1 & "artian visitor0 notin! this0 mi!ht infer that St &(s chime
ca"sed St B(s to chime1 We0 of course0 kno% etter0 ut the only real test of the hypothesis %ould e
experimentally to rin! the St &(s chime at random times rather than once per hour1 The "artian(s
prediction 7%hich %ould of course e disproved in this case8 is that St B(s clock %ill still chime
immediately after St &(s1 #t is only experimental manipulation that can determine %hether an
oserved correlation truly indicates causation1
#f your hypothesis is that the non:random survival of random !enetic variation has important
evolutionary conseEuences0 the experimental test of the hypothesis %ould have to e a delierate
human intervention1 Go in and manip"late %hich variant survives and %hich doesn(t1 Go in there
and choose0 as a human reeder0 %hich kinds of individuals !et to reproduce1 &nd that0 of course0 is
artificial selection1 &rtificial selection is not Hust an analo$y for natural selection1 &rtificial
selection constitutes a true experimental : as opposed to oservational : test of the hypothesis that
selection causes evolutionary chan!e1
"ost of the kno%n examples of artificial selection : for example0 the manufacture of the
various reeds of do! : are oserved %ith the hindsi!ht of history0 rather than ein! delierate tests
of predictions under experimentally controlled conditions1 But proper experiments have een done0
and the results have al%ays een as expected from the more anecdotal results on do!s0 caa!es and
sunflo%ers1 )ere is a typical example0 an especially !ood one ecause a!ronomists at the #llinois
Experimental Station e!an the experiment rather a lon! time a!o0 in 2;9< 7Generation 2 in the
!raph81 The dia!ram aove sho%s the oil content in maiFe seeds of t%o different artificially selected
lines0 one selected for hi!h oil yield0 and the other for lo% oil yield1 This is a true experiment
ecause %e are comparin! the results of t%o delierate manipulations or interventions1 Evidently
the difference is dramatic0 and it increases1 #t seems likely that oth the up%ard trend and the
do%n%ard trend %ould eventually level offA the lo%:yieldin! line ecause you can(t drop elo%
Fero oil content0 and the hi!h:yieldin! line for reasons that are nearly as plain1
T,o lines o$ "ai-e selete% $or hi'h an% lo, oil ontent
)ere(s a further laoratory demonstration of the po%er of artificial selection0 %hich is
instructive in another %ay1 The dia!ram overleaf sho%s some seventeen !enerations of rats0
artificially selected for resistance to tooth decay1 The measure ein! plotted is the time0 in days0 that
the rats %ere free of dental caries1 &t the start of the experiment0 the typical period free of tooth
decay %as aout 255 days1 &fter only a doFen or so !enerations of systematic selection a!ainst
caries0 the decay:free period %as aout four times as lon!0 or even more1 Once a!ain0 a separate line
%as selected to evolve in the opposite directionA in this case the experiments systematically red for
susceptiility to tooth decay1
T,o lines o$ rats selete% $or hi'h an% lo, resistane to tooth %eay
The example offers an opportunity to cut our teeth on natural selection thinkin!1 #ndeed0 this
discussion of rat teeth %ill e the first of three such excursions into natural selection proper0 %hich
%e are no% eEuipped to undertake1 #n the other t%o0 as %ith the rats0 %e shall revisit creatures
already met alon! the (primrose path( from domestication0 namely do!s and flo%ers1
Why0 if it is so easy to improve the teeth of rats y artificial selection0 did natural selection
apparently make such a poor Ho of it in the first placeB Surely there is no enefit in tooth decay1
Why0 if artificial selection is capale of reducin! it0 didn(t natural selection do the same Ho lon!
a!oB # can think of t%o ans%ers0 oth instructive1
The first ans%er is that the ori!inal population that the human selectors used as their ra%
material consisted not of %ild rats ut of domesticated laoratory:red %hite rats1 #t could e said
that la rats are feather:edded0 like modern humans0 shielded from the cuttin! ed!e of natural
selection1 & !enetic tendency to tooth decay %ould si!nificantly reduce reproductive prospects in
the %ild0 ut mi!ht make no difference in a laoratory colony %here the livin! is easy0 and the
decision on %ho reeds and %ho does not is taken y humans0 %ith no eye to survival1
That(s the first ans%er to the Euestion1 The second ans%er is more interestin!0 for it carries
an important lesson aout natural selection0 as %ell as artificial selection1 #t is the lesson of trade:
offs0 and %e have already adverted to it %hen talkin! aout pollination strate!ies in plants1 *othin!
is free0 everythin! comes %ith a price ta!1 #t mi!ht seem ovious that tooth decay is to e avoided at
all costs0 and # do not dout that dental caries si!nificantly shortens life in rats1 But let(s think for a
moment aout %hat must happen in order to increase an animal(s resistance to tooth decay1 # don(t
kno% the details0 ut # am confident that it %ill e costly0 and that is all # need to assume1 -et us
suppose it is achieved y a thickenin! of the %all of the tooth0 and this reEuires extra calcium1 #t is
not impossile to find extra calcium0 ut it has to come from some%here0 and it is not free1 Calcium
7or %hatever the limitin! resource mi!ht e8 is not floatin! around in the air1 #t has to come into the
ody via food1 &nd it is potentially useful for other thin!s apart from teeth1 The ody has somethin!
%e could call a calcium economy1 Calcium is needed in one0 and it is needed in milk1 7#(m
assumin! it is calcium %e are talkin! aout1 Even if it is not calcium0 there must e some costly
limitin! resource0 and the ar!ument %ill %ork Hust as %ell0 %hatever the limitin! resource is1 #(ll
continue to use calcium for the sake of ar!ument18 &n individual rat %ith extra stron! teeth mi!ht
%ell tend to live lon!er than a rat %ith rotten teeth0 all other thin!s ein! eEual1 But all other thin!s
are not eEual0 ecause the calcium needed to stren!then the teeth had to come from some%here0 say0
ones1 & rival individual %hose !enes did not predispose it to take calcium a%ay from ones mi!ht
conseEuently survive lon!er0 ecause of its superior ones and in spite of its ad teeth1 Or the rival
individual mi!ht e etter Eualified to rear children ecause she makes more calcium:rich milk1 &s
economists are fond of Euotin! from Roert )einlein0 there(s no such thin! as a free lunch1 "y rat
example is hypothetical0 ut it is safe to say that0 for economic reasons0 there must e such a thin!
as a rat %hose teeth are too perfect1 Perfection in one department must e ou!ht0 in the form of a
sacrifice in another department1
The lesson applies to all livin! creatures1 We can expect odies to e %ell eEuipped to
survive0 ut this does not mean they should e perfect %ith respect to any one dimension1 &n
antelope mi!ht run faster0 and e more likely to escape a leopard0 if its le!s %ere a little lon!er1 But
a rival antelope %ith lon!er le!s0 althou!h it mi!ht e etter eEuipped to outsprint a predator0 has to
pay for its lon! le!s in some other department of the ody(s economy1 The materials needed to
make the extra one and muscle in the lon!er le!s have to e taken from some%here else0 so the
lon!er:le!!ed individual is more likely to die for reasons other than predation1 Or it may even e
more likely to die from predation ecause its lon!er le!s0 althou!h they can run faster %hen intact0
are more likely to reak0 in %hich case it can(t run at all1 & ody is a patch%ork of compromises1 #
shall return to this point in the chapter on arms races1
What happens under domestication is that animals are artificially shielded from many of the
risks that shorten the lives of %ild animals1 & pedi!ree dairy co% may yield prodi!ious Euantities of
milk0 ut its pendulously cumersome udder %ould seriously impede it in any attempt to outrun a
lion1 Thorou!hred horses are super runners and Humpers0 ut their le!s are vulnerale to inHury
durin! races0 especially over Humps0 %hich su!!ests that artificial selection has pushed them into a
Fone that natural selection %ould not have tolerated1 "oreover0 Thorou!hreds thrive only on a rich
diet supplied y humans1 Whereas Britain(s native ponies0 for example0 flourish on pasture0
racehorses don(t prosper unless they are fed a much richer diet of !rains and supplements : %hich
they %ould not find in the %ild1 &!ain0 #(ll return to such matters in the arms race chapter1
'OGS &G&#*
)avin! finally reached the topic of natural selection0 %e can turn ack to the example of
do!s for some other important lessons1 # said that they are domesticated %olves0 ut # need to
Eualify this in the li!ht of a fascinatin! theory of the evolution of the do!0 %hich has a!ain een
most clearly articulated y Raymond Coppin!er1 The idea is that the evolution of the do! %as not
Hust a matter of artificial selection1 #t %as at least as much a case of %olves adaptin! to the %ays of
man y natural selection1 "uch of the initial domestication of the do! %as self0 domestication0
mediated y natural0 not artificial0 selection1 -on! efore %e !ot our hands on the chisels in the
artificial selection toolox0 natural selection had already sculpted %olves into self:domesticated
(villa!e do!s( %ithout any human intervention1 Only later did humans adopt these villa!e do!s and
transmo!rify them0 separately and comprehensively0 into the raino% spectrum of reeds that today
!race 7if !race is the %ord8 Crufts and similar pa!eants of canine achievement and eauty 7if eauty
is the %ord81
Coppin!er points out that %hen domestic animals reak free and !o feral for many
!enerations0 they usually revert to somethin! close to their %ild ancestor1 We mi!ht expect feral
do!s0 therefore0 to ecome rather %olf:like1 But this doesn(t happen1 #nstead0 do!s left to !o feral
seem to ecome the uiEuitous (villa!e do!s( : (pye:do!s( : that han! around human settlements all
over the third %orld1 This encoura!es Coppin!er(s elief that the do!s on %hich human reeders
finally %ent to %ork %ere %olves no lon!er1 They had already chan!ed themselves into do!sA
villa!e do!s0 pye:do!s0 perhaps din!os1
Real %olves are pack hunters1 +illa!e do!s are scaven!ers that freEuent middens and
ruish dumps1 Wolves scaven!e too0 ut they are not temperamentally suited to scaven!in! human
ruish ecause of their lon! (fli!ht distance(1 #f you see an animal feedin!0 you can measure its
fli!ht distance y seein! ho% close it %ill let you approach efore fleein!1 ,or any !iven species in
any !iven situation0 there %ill e an optimal fli!ht distance0 some%here et%een too risky or
foolhardy at the short end0 and too fli!hty or risk:averse at the lon! end1 #ndividuals that take off
too late %hen dan!er threatens are more likely to e killed y that very dan!er1 -ess oviously0
there is such a thin! as takin! off too soon1 #ndividuals that are too fli!hty never !et a sEuare meal0
ecause they run a%ay at the first hint of dan!er on the horiFon1 #t is easy for us to overlook the
dan!ers of ein! too risk:averse1 We are puFFled %hen %e see Feras or antelopes calmly !raFin! in
full vie% of lions0 keepin! no more than a %ary eye on them1 We are puFFled0 ecause our o%n risk
aversion 7or that of our safari !uide8 keeps us firmly inside the -and Rover even thou!h %e have no
reason to think there is a lion %ithin miles1 This is ecause %e have nothin! to set a!ainst our fear1
We are !oin! to !et our sEuare meals ack at the safari lod!e1 Our %ild ancestors %ould have had
much more sympathy %ith the risk:takin! Feras1 -ike the Feras0 they had to alance the risk of
ein! eaten a!ainst the risk of not eatin!1 Sure0 the lion mi!ht attackJ ut0 dependin! on the siFe of
your troop0 the odds %ere that it %ould catch another memer of it rather than you1 &nd if you
never ventured on to the feedin! !rounds0 or do%n to the %ater hole0 you(d die any%ay0 of hun!er or
thirst1 #t is the same lesson of economic trade:offs that %e have already met0 t%ice1 L
The ottom line of that di!ression is that the %ild %olf0 like any other animal0 %ill have an
optimal fli!ht distance0 nicely poised : and potentially flexile : et%een too old and too fli!hty1
*atural selection %ill %ork on the fli!ht distance0 movin! it one %ay or the other alon! the
continuum if conditions chan!e over evolutionary time1 #f a plenteous ne% food source in the form
of villa!e ruish dumps enters the %orld of %olves0 that is !oin! to shift the optimum point
to%ards the shorter end of the fli!ht distance continuum0 in the direction of reluctance to flee %hen
enHoyin! this ne% ounty1
We can ima!ine %ild %olves scaven!in! on a ruish tip on the ed!e of a villa!e1 "ost of
them0 fearful of men thro%in! stones and spears0 have a very lon! fli!ht distance1 They sprint for
the safety of the forest as soon as a human appears in the distance1 But a fe% individuals0 y !enetic
chance0 happen to have a sli!htly shorter fli!ht distance than the avera!e1 Their readiness to take
sli!ht risks : they are rave0 shall %e say0 ut not foolhardy : !ains them more food than their more
risk:averse rivals1 &s the !enerations !o y0 natural selection favours a shorter and shorter fli!ht
distance0 until Hust efore it reaches the point %here the %olves really are endan!ered y stone:
thro%in! humans1 The optimum fli!ht distance has shifted ecause of the ne%ly availale food
Somethin! like this evolutionary shortenin! of the fli!ht distance %as0 in Coppin!er(s vie%0
the first step in the domestication of the do!0 and it %as achieved y natural selection0 not artificial
selection1 'ecreasin! fli!ht distance is a ehavioural measure of %hat mi!ht e called increasin!
tameness1 &t this sta!e in the process0 humans %ere not delierately choosin! the tamest individuals
for reedin!1 &t this early sta!e0 the only interactions et%een humans and these incipient do!s
%ere hostile1 #f %olves %ere ecomin! domesticated it %as y self:domestication0 not delierate
domestication y people1 'elierate domestication came later1
We can !et an idea of ho% tameness0 or anythin! else0 can e sculpted : naturally or
artificially : y lookin! at a fascinatin! experiment of modern times0 on the domestication of
Russian silver foxes for use in the fur trade1 #t is douly interestin! ecause of the lessons it teaches
us0 over and aove %hat 'ar%in kne%0 aout the domestication process0 aout the (side:effects( of
selective reedin!0 and aout the resemlance0 %hich 'ar%in %ell understood0 et%een artificial
and natural selection1
The silver fox is Hust a colour variant0 valued for its eautiful fur0 of the familiar red fox0
1"lpes "lpes1 The Russian !eneticist 'imitri Belyaev %as employed to run a fox fur farm in the
29=5s1 )e %as later sacked ecause his scientific !enetics conflicted %ith the anti:scientific
ideolo!y of -ysenko0 the charlatan iolo!ist %ho mana!ed to capture the ear of Stalin and so take
over0 and lar!ely ruin0 all of Soviet !enetics and a!riculture for some t%enty years1 Belyaev retained
his love of foxes0 and of true -ysenko:free !enetics0 and he %as later ale to resume his studies of
oth0 as director of an #nstitute of Genetics in Sieria1
Wild foxes are tricky to handle0 and Belyaev set out delierately to reed for tameness1 -ike
any other animal or plant reeder of his time0 his method %as to exploit natural variation 7no !enetic
en!ineerin! in those days8 and choose0 for reedin!0 those males and females that came closest to
the ideal he %as seekin!1 #n selectin! for tameness0 Belyaev could have chosen0 for reedin!0 those
do!s and itches that most appealed to him0 or looked at him %ith the cutest facial expressions1 That
mi!ht %ell have had the desired effect on the tameness of future !enerations1 "ore systematically
than that0 ho%ever0 he used a measure that %as pretty close to the (fli!ht distance( # Hust mentioned
in connection %ith %ild %olves0 ut adapted for cus1 Belyaev and his collea!ues 7and successors0
for the experimental pro!ram continued after his death8 suHected fox cus to standardiFed tests in
%hich an experimenter %ould offer a cu food y hand0 %hile tryin! to stroke or fondle it1 The cus
%ere classified into three classes1 Class ### cus %ere those that fled from or it the person1 Class ##
cus %ould allo% themselves to e handled0 ut sho%ed no positive responsiveness to the
experimenters1 Class # cus0 the tamest of all0 positively approached the handlers0 %a!!in! their
tails and %hinin!1 When the cus !re% up0 the experimenters systematically red only from this
tamest class1
&fter a mere six !enerations of this selective reedin! for tameness0 the foxes had chan!ed
so much that the experimenters felt oli!ed to name a ne% cate!ory0 the (domesticated elite( class0
%hich %ere (ea!er to estalish human contact0 %himperin! to attract attention and sniffin! and
lickin! experimenters like do!s(1 &t the e!innin! of the experiment0 none of the foxes %ere in the
elite class1 &fter ten !enerations of reedin! for tameness0 2; per cent %ere (elite(J after t%enty
!enerations0 4= per centJ and after thirty to thirty:five !enerations0 (domesticated elite( individuals
constituted et%een @5 and ;5 per cent of the experimental population1
Such results are perhaps not too surprisin!0 except for the astonishin! ma!nitude and speed
of the effect1 Thirty:five !enerations %ould pass unnoticed on the !eolo!ical timescale1 Even more
interestin!0 ho%ever0 %ere the unexpected side:effects of the selective reedin! for tameness1 These
%ere truly fascinatin! and !enuinely unforeseen1 'ar%in0 the do!:lover0 %ould have een
entranced1 The tame foxes not only ehaved like domestic do!s0 they looked like them1 They lost
their foxy pela!e and ecame pieald lack and %hite0 like Welsh collies1 Their foxy prick ears
%ere replaced y do!!y floppy ears1 Their tails turned up at the end like a do!(s0 rather than do%n
like a fox(s rush1 The females came on heat every six months like a itch0 instead of every year like
a vixen1 &ccordin! to Belyaev0 they even sounded like do!s1
!elaye. ,ith his $o/es, as they t(rn ta"e ) an% %o'li&e
These do!:like features %ere side:effects1 Belyaev and his team did not delierately reed
for them0 only for tameness1 Those other do!:like characteristics seemin!ly rode on the
evolutionary coattails of the !enes for tameness1 To !eneticists0 this is not surprisin!1 They
reco!niFe a %idespread phenomenon called (pleiotropy(0 %herey !enes have more than one effect0
seemin!ly unconnected1 The stress is on the %ord (seemin!ly(1 Emryonic development is a
complicated usiness1 &s %e learn more aout the details0 that (seemin!ly unconnected( turns into
(connected y a route that %e no% understand0 ut didn(t efore(1 Presumaly !enes for floppy ears
and pieald coats are pleiotropically linked to !enes for tameness0 in foxes as %ell as in do!s1 This
illustrates a !enerally important point aout evolution1 When you notice a characteristic of an
animal and ask %hat its 'ar%inian survival value is0 you may e askin! the %ron! Euestion1 #t
could e that the characteristic you have picked out is not the one that matters1 #t may have (come
alon! for the ride(0 dra!!ed alon! in evolution y some other characteristic to %hich it is
pleiotropically linked1
The evolution of the do!0 then0 if Coppin!er is ri!ht0 %as not Hust a matter of artificial
selection0 ut a complicated mixture of natural selection 7%hich predominated in the early sta!es of
domestication8 and artificial selection 7%hich came to the fore more recently81 The transition %ould
have een seamless0 %hich a!ain !oes to emphasiFe the similarity : as 'ar%in reco!niFed : et%een
artificial and natural selection1
,-OWERS &G&#*
-et(s no%0 in the third of our %arm:up forays into natural selection0 move on to flo%ers and
pollinators and see somethin! of the po%er of natural selection to drive evolution1 Pollination
iolo!y furnishes us %ith some pretty amaFin! facts0 and the hi!h point of %ondrousness is reached
in the orchids1 *o %onder 'ar%in %as so keen on themJ no %onder he %rote the ook # have
already mentioned0 The 1ario"s #ontriances by which !rchids are ,ertilised by &nsects1 Some
orchids0 such as the (ma!ic ullet( "ada!ascar ones %e met earlier0 !ive nectar0 ut others have
found a %ay to ypass the costs of feedin! pollinators0 y trickin! them instead1 There are orchids
that resemle female ees 7or %asps or flies8 %ell enou!h to fool males into attemptin! to copulate
%ith them1 To the extent that such mimics resemle females of one particular insect species0 to that
extent %ill males of those species serve as ma!ic ullets0 !oin! from flo%er to flo%er of Hust the one
orchid species1 Even if the orchid resemles (any old ee( rather than one species of ee0 the ees
that it fools %ill still e (fairly ma!ic( ullets1 #f you or # %ere to look closely at a fly orchid or a ee
orchid 7see colour pa!e =80 %e %ould e ale to tell that it %as not a real insectJ ut %e %ould e
fooled at a casual !lance out of the corner of our eye1 &nd even lookin! at it head:on0 # %ould say
the ee orchid in the picture 7h8 is pretty clearly more of a umle:ee orchid than a honey:ee
orchid1 #nsects have compound eyes0 %hich are not so acute as our camera eyes0 and the shapes and
colours of insect:mimickin! orchids0 reinforced y seductive scents that mimic those of female
insects0 are more than capale of trickin! males1 By the %ay0 it is Euite proale that the mimicry is
enhanced %hen seen in the ultraviolet ran!e0 from %hich %e are cut off1
The so:called spider orchid0 Brassia 7colour pa!e = 7k880 achieves pollination y a different
kind of deception1 The females of various species of solitary %asp 7(solitary( ecause they don(t live
socially in lar!e nests like the familiar autumn pests0 called yello%Hackets y &mericans8 capture
spiders0 stin! them to paralyse them0 and lay their e!!s on them as a livin! food supply for their
larvae1 Spider orchids resemle spiders sufficiently to fool female %asps into attemptin! to stin!
them1 #n the process they pick up pollinia : masses of pollen !rains produced y the orchids1 When
they move on to try to stin! another spider orchid0 the pollinia are transferred1 By the %ay0 # can(t
resist addin! the exactly ack%ards case of the spider Epicad"s hetero$aster0 %hich mimics an
orchid1 #nsects come to the (flo%er( in search of nectar0 and are promptly eaten y it1
Some of the most astonishin! orchids that practise this seduction trick are to e found in
Western &ustralia1 +arious species in the !enus Drakaea are kno%n as hammer orchids1 Each
species has a special relationship %ith a particular species of %asp of the type called thynnids1 Part
of the flo%er ears a crude resemlance to an insect0 dupin! the male thynnid %asp into attemptin!
to mate %ith it1 So far in my description0 Drakaea is not dramatically different from other insect:
mimickin! orchids1 But Drakaea has a remarkale extra trick up its sleeveA the fake (%asp( is orne
on the end of a hin!ed (arm(0 %ith a flexile (elo%(1 6ou can clearly see the hin!e in the picture
7colour pa!e = 7!881 The flutterin! movement of the %asp !rippin! the dummy %asp causes the
(elo%( to end0 and the %asp is dashed repeatedly ack and forth like a hammer a!ainst the other
side of the flo%er : let(s call it the anvil : %here it keeps its sexual parts1 The pollinia are dislod!ed
and stick to the %asp0 %ho eventually extricates himself and flies off0 sadder ut apparently no
%iserA he !oes on to repeat the performance on another hammer orchid0 %here he and the pollinia he
ears are duly dashed a!ainst the anvil0 so that his car!o finds its destined refu!e on the female
or!ans of the flo%er1 # sho%ed a film of this astoundin! performance in one of my Royal #nstitution
Christmas -ectures for Children0 and it can e seen in the recordin! of the lecture called (The
$ltraviolet Garden(1
#n the same lecture # discussed the (ucket orchids( of South &merica0 %hich achieve
pollination in an eEually remarkale ut rather different %ay1 They too have specialiFed pollinators0
not %asps ut small ees0 of the !roup called Eu!lossine1 &!ain0 these orchids provide no nectar1
But the orchids don(t fool the ees into matin! %ith them either1 #nstead0 they provide a vital piece
of assistance for male ees0 %ithout %hich the ees %ould e unale to attract real females1
These little ees0 %hich live only in South &merica0 have a stran!e hait1 They !o to
elaorate len!ths to collect fra!rant0 or any%ay smelly0 sustances0 %hich they store in special
containers attached to their enlar!ed hind le!s1 #n different species these smelly sustances can
come from flo%ers0 from dead %ood0 or even from faeces1 #t seems that they use the !athered
perfumes to attract0 or other%ise court0 females1 "any insects use particular scents to appeal to the
opposite sex0 and most of them manufacture the perfumes in special !lands1 ,emale silk moths0 for
example0 attract males from an astonishin!ly lon! distance y releasin! a uniEue scent0 %hich they
manufacture and %hich males detect : in minute traces from literally miles a%ay : %ith their
antennae1 #n the case of Eu!lossine ees0 it is the males that use scent1 &nd0 unlike the female
moths0 they don(t synthesiFe their o%n perfume ut use the smelly in!redients that they have
collected0 not as pure sustances ut as carefully concocted lends %hich they put to!ether like
expert perfumiers1 Each species mixes a characteristic cocktail of sustances !athered from various
sources1 &nd there are some species of Eu!lossine ee that positively need0 for manufacturin! their
characteristic species scent0 sustances that are supplied only y flo%ers of particular species of the
orchid !enus #oryanthes 0 ucket orchids1 The common name of Eu!lossine ees is (orchid ees(1
What an intricate picture of mutual dependence1 The orchids need the Eu!lossine ees0 for
the usual (ma!ic ullet( reasons1 &nd the ees need the orchids0 for the rather %eirder reason that
they can(t attract female ees %ithout sustances that are either impossile or at least too hard to
find except throu!h the !ood offices of ucket orchids1 But the %ay in %hich pollination is achieved
is even %eirder still0 and it superficially makes the ee look more like a victim than a cooperatin!
& male Eu!lossine ee is attracted to the orchids y the smell of the sustances that he
needs in order to manufacture his sexual perfumes1 )e ali!hts on the rim of the ucket and starts to
scrape the %axy perfume into the special scent pockets in his le!s1 But the rim of the ucket is
slippery underfoot : and there(s a reason for this1 The ee falls into the ucket0 %hich is filled %ith
liEuid0 in %hich he s%ims1 )e cannot clim up the slippery sides of the ucket1 There is only one
escape route0 and this is a special ee:siFed hole in the side of the ucket 7not visile in the picture
that appears on colour pa!e >81 )e is !uided y (steppin! stones( to the hole and starts to cra%l
throu!h it1 #t(s a ti!ht fit0 and it ecomes even ti!hter as the (Ha%s( 7these you can see in the pictureA
they look like the chuck of a lathe or electric drill8 contract and trap him1 While he is held in their
!rip0 they !lue t%o pollinia to his ack1 The !lue takes a %hile to set0 after %hich the Ha%s a!ain
relax and release the ee0 %ho flies off0 complete %ith pollinia on his ack1 Still in search of the
precious in!redients for his perfumery0 the ee lands on another ucket orchid and the process
repeats itself1 This time0 ho%ever0 as the ee stru!!les throu!h the hole in the ucket0 the pollinia
are scraped off0 and they fertiliFe the sti!ma of this second orchid1
The intimate relationship et%een flo%ers and their pollinators is a lovely example of %hat
is called co:evolution : evolution to!ether1 Co:evolution often occurs et%een or!anisms that have
somethin! to !ain from each other0 partnerships in %hich each side contriutes somethin! to the
other0 and oth !ain from the cooperation1 &nother eautiful example is the set of relationships that
have !ro%n up around coral reefs0 independently in many different parts of the %orld0 et%een
cleaner fish and lar!er fish1 The cleaners elon! to several different species0 and some are not even
fish at all ut shrimps : a nice case of conver!ent evolution1 Cleanin!0 amon! coral:reef fish0 is a
%ell:estalished %ay of life0 like huntin! or !raFin! or anteatin! amon! mammals1 Cleaners make
their livin! y pickin! parasites off the odies of their lar!er (clients(1 That the clients enefit has
een ele!antly demonstrated y removin! all the cleaners from an experimental area of reef0
%hereupon the health of lots of species of fish declines1 # have discussed the cleanin! hait
else%here0 so %ill say no more here1
Co:evolution also occurs et%een species that don(t enefit from each other(s presence0 like
predators and prey0 or parasites and hosts1 These kinds of co:evolution are sometimes called (arms
races( and # postpone discussin! them to Chapter 231
-et me dra% this chapter0 and the previous one0 to a conclusion1 Selection : in the form of
artificial selection y human reeders : can turn a pye:do! into a Pekinese0 or a %ild caa!e into a
cauliflo%er0 in a fe% centuries1 The difference et%een any t%o reeds of do! !ives us a rou!h idea
of the Euantity of evolutionary chan!e that can e achieved in less than a millennium1 The next
Euestion %e should ask is0 ho% many millennia do %e have availale to us in accountin! for the
%hole history of lifeB #f %e ima!ine the sheer Euantity of difference that separates a pye:do! from a
peke0 %hich took only a fe% centuries of evolution0 ho% much lon!er is the time that separates us
from the e!innin! of evolution or0 say0 from the e!innin! of the mammalsB Or from the time
%hen fish emer!ed on to the landB The ans%er is that life e!an not Hust centuries a!o ut tens of
millions of centuries a!o1 The measured a!e of our planet is aout >1< illion years0 or aout ><
million centuries1 The time that has elapsed since the common ancestor of all today(s mammals
%alked the Earth is aout t%o million centuries1 & century seems a pretty lon! time to us1 Can you
ima!ine t%o million centuries0 laid end to endB The time that has elapsed since our fish ancestors
cra%led out of the %ater on to the land is aout three and a half million centuriesA that is to say0
aout t%enty thousand times as lon! as it took to make all the different : really very different :
reeds of do!s from the common ancestor that they all share1
)old in your head an approximate picture of the Euantity of difference et%een a peke and a
pye:do!1 We aren(t talkin! precise measurements hereA it %ould do Hust as %ell to think aout the
difference et%een any one reed of do! and any other0 for that is on avera!e doule the amount of
chan!e that has een %rou!ht0 y artificial selection0 from the common ancestor1 Bear in mind this
order of evolutionary chan!e0 and then extrapolate ack%ards t%enty thousand times as far into the
past1 #t ecomes rather easy to accept that evolution could accomplish the amount of chan!e that it
took to transform a fish into a human1
But all this presupposes that %e kno% the a!e of the Earth0 and of the various landmark
points in the fossil record1 This is a ook aout evidence0 so # can(t Hust assert dates ut must Hustify
them1 )o%0 actually0 do %e kno% the a!e of any particular rockB )o% do %e kno% the a!e of a
fossilB )o% do %e kno% the a!e of the EarthB )o%0 for that matter0 do %e kno% the a!e of the
universeB We need clocks0 and clocks are the suHect of the next chapter1
L &s in all memers of the daisy family0 each (flo%er( is actually many little flo%ers
7florets80 undled to!ether in the dark disk in the middle1 The yello% petals that surround the
sunflo%er are in fact the petals of Hust the florets around the ed!e1 The florets in the rest of the disk
have petals0 ut too small to e noticed1
O Perhaps ecause : ein! a *e% World plant : the sunflo%er is not mentioned explicitly in
the Bile1 The theolo!ical mind takes a deli!ht in the niceties of dietary la%s and the in!enuity
reEuired to dod!e them1 #n South &merica0 capyaras 7sort of !iant !uinea pi!s8 %ere deemed to e
honorary fish for the purposes of Catholic dietary la%s on ,ridays0 presumaly ecause they live in
%ater1 &ccordin! to the food %riter 'oris Reynolds0 ,rench Catholic !ourmets discovered a
loophole that enaled them to eat meat on ,ridays1 -o%er a le! of lam into a %ell and then (fish( it
out1 They must think God is a%fully easily fooled1
L Oliver "orton discusses this and related issues in his provokin!ly lyrical ook Eatin$ the
L &t least there is no reason to think that they do0 or indeed that they enHoy anythin! in the
sense %e understand1 # shall return to this perennial temptation in Chapter 231
L Which %ere in turn named after the (multitude of do!s of a hu!e siFe( mentioned in Pliny(s
.at"ral )istory1
L ,or example0 herdin! in sheepdo!s is derived from stalkin! in %olves0 %ith the killin!
removed from the end of the seEuence1
L #t doesn(t affect the point # am makin!0 ut this story applies only to female an!ler fish1
The males are usually tiny d%arfs0 %ho attach themselves parasitically to a female(s ody0 like a
little extra fin1
L The popular canard aout )itler ein! inspired y 'ar%in comes partly from the fact that
oth )itler and 'ar%in %ere impressed y somethin! that everyody has kno%n for centuriesA you
can reed animals for desired Eualities1 )itler aspired to turn this common kno%led!e to the human
species1 'ar%in didn(t1 )is inspiration took him in a much more interestin! and ori!inal direction1
'ar%in(s !reat insi!ht %as that you don(t need a reedin! a!ent at allA nature : ra% survival or
differential reproductive success : can play the role of reeder1 &s for )itler(s ( Social 'ar%inism( :
his elief in a stru!!le et%een races : that is actually very "n:'ar%inian1 ,or 'ar%in0 the stru!!le
for existence %as a stru!!le et%een individuals %ithin a species0 not et%een species0 races or
other !roups1 'on(t e misled y the ill:chosen and unfortunate sutitle of 'ar%in(s !reat ookA The
preseration of fao"red races in the str"$$le for life1 #t is aundantly clear from the text itself that
'ar%in didn(t mean races in the sense of (& !roup of people0 animals0 or plants0 connected y
common descent or ori!in( 7 !xford En$lish Dictionary0 definition <1#81 Rather0 he intended
somethin! more like the !ED(s definition <1##A (& !roup or class of people0 animals0 or thin!s0
havin! some common feature or features(1 &n example of sense <1 ## %ould e (&ll those individuals
7re!ardless of their !eo!raphical race8 %ho have lue eyes(1 #n the technical Har!on of modern
!enetics0 %hich %as not availale to 'ar%in0 %e %ould express the sense of (race( in his sutitle as
(&ll those individuals %ho possess a certain allele1( The misunderstandin! of the 'ar%inian stru!!le
for existence as a stru!!le et%een $ro"ps of individuals : the so:called (!roup selection( fallacy : is
unfortunately not confined to )itlerian racism1 #t constantly resurfaces in amateur misinterpretations
of 'ar%inism0 and even amon! some professional iolo!ists %ho should kno% etter1
L Psycholo!ists have analo!ous tests of risk:takin! amon! humans0 %hich sho% interestin!
differences1 Entrepreneurs typically score hi!hly on risk:takin! measures0 as do pilots0 rock:
climers0 motorcycle racers and other extreme sports enthusiasts1 Women tend to e more risk:
averse than men1 ,eminists %ill here point out that the causal arro% could !o either %ayA %omen
could ecome more risk:averse ecause of the occupations into %hich society thrusts them1
S#-E*CE &*' S-OW T#"E
# , the history:deniers %ho dout the fact of evolution are i!norant of iolo!y0 those %ho
think the %orld e!an less than ten thousand years a!o are %orse than i!norant0 they are deluded to
the point of perversity1 They are denyin! not only the facts of iolo!y ut those of physics0 !eolo!y0
cosmolo!y0 archaeolo!y0 history and chemistry as %ell1 This chapter is aout ho% %e kno% the
a!es of rocks and the fossils emedded in them1 #t presents the evidence that the timescale on %hich
life has operated on this planet is measured not in thousands of years ut in thousands of millions of
Rememer0 evolutionary scientists are in the position of detectives %ho come late to the
scene of a crime1 To pinpoint %hen thin!s happened0 %e depend upon traces left y time:dependent
processes : clocks0 in a road sense1 One of the first thin!s a detective does %hen investi!atin! a
murder is ask a doctor or patholo!ist to estimate the time of death1 "uch follo%s from this
information0 and in detective fiction an almost mystical reverence is accorded to the patholo!ist(s
estimate1 The (time of death( is a aseline fact0 an inerrant pivot around %hich more or less far:
fetched speculations y the detective revolve1 But that estimate is0 of course0 suHect to error0 an
error that can e measured and can e Euite lar!e1 The patholo!ist uses various time:dependent
processes to estimate the time of deathA the ody cools at a characteristic rate0 ri!or mortis sets in at
a particular time0 and so on1 These are the rather crude (clocks( availale to the investi!ator of a
murder1 The clocks availale to the evolutionary scientist are potentially much more accurate : in
proportion to the timescale involved0 of course0 not more accurate to the nearest hourK The analo!y
to a precision clock is more persuasive for a .urassic rock in the hands of a !eolo!ist than it is for a
coolin! corpse in the hands of a patholo!ist1
"an:made clocks %ork on timescales that are very short y evolutionary standards : hours0
minutes0 seconds : and the time:dependent processes they use are fastA the s%in!in! of a pendulum0
the s%ivellin! of a hairsprin!0 the oscillation of a crystal0 the urnin! of a candle0 the drainin! of a
%ater vessel or an hour!lass0 the rotation of the earth 7re!istered y a sundial81 &ll clocks exploit
some process that occurs at a steady and kno%n rate1 & pendulum s%in!s at a very constant rate0
%hich depends upon its len!th ut not0 at least in theory0 on the amplitude of the s%in! or the mass
of the o on the end1 Grandfather clocks %ork y linkin! a pendulum to an escapement %hich
advances a toothed %heel0 step y stepJ the rotation is then !eared do%n to the speed of rotation of
an hour hand0 a minute hand and a second hand1 Watches %ith hairsprin! %heels %ork in a similar
%ay1 'i!ital %atches exploit an electronic eEuivalent of a pendulum0 the oscillation of certain kinds
of crystals %hen supplied %ith ener!y from a attery1 Water clocks and candle clocks are much less
accurate0 ut they %ere useful efore the invention of event:countin! clocks1 They depend not on
countin! thin!s0 as a pendulum clock or a di!ital %atch does0 ut on measurin! some Euantity1
Sundials are inaccurate %ays of tellin! the time1L But the rotation of the earth0 %hich is the time:
dependent process on %hich they rely0 is accurate on the timescale of the slo%er clock that %e call
the calendar1 This is ecause on that timescale it is no lon!er a measurin! clock 7a sundial measures
the continuously varyin! an!le of the sun8 ut a countin! clock 7countin! dayGni!ht cycles81
Both countin! clocks and measurin! clocks are availale to us on the immensely slo%
timescale of evolution1 But to investi!ate evolution %e don(t need Hust a clock that tells the present
time0 as a sundial does0 or a %atch1 We need somethin! more like a stop%atch that can e reset1 Our
evolutionary clock needs to e 2eroed at some point0 so that %e can calculate the elapsed time since
a startin! point0 to !ive us0 for example0 the asolute a!e of some oHect such as a rock1 Radioactive
clocks for datin! i!neous 7volcanic8 rocks are conveniently Feroed at the moment the rock is formed
y the solidification of molten lava1
,ortunately0 a variety of Fero:ale natural clocks is availale1 This variety is a !ood thin!0
ecause %e can use some clocks to check the accuracy of other clocks1 Even more fortunately0 they
sensitively cover an astonishin!ly %ide ran!e of timescales0 and %e need this too ecause
evolutionary timescales span seven or ei!ht orders of ma!nitude1 #t(s %orth spellin! out %hat this
means1 &n order of ma!nitude means somethin! precise1 & chan!e of one order of ma!nitude is one
multiplication 7or division8 y ten1 Since %e use a decimal system0L the order of ma!nitude of a
numer is a count of the numer of Feroes0 efore or after the decimal point1 So a ran!e of ei!ht
orders of ma!nitude constitutes a hundred millionfold1 The second hand of a %atch rotates <5 times
as fast as the minute hand and @35 times as fast as the hour hand0 so the three hands cover a ran!e
%hich is less than three orders of ma!nitude1 This is tiny compared to the ei!ht orders of ma!nitude
spanned y our repertoire of !eolo!ical clocks1 Radioactive decay clocks are availale for short
timescales as %ell0 even do%n to fractions of a secondJ ut for evolutionary purposes0 clocks that
can measure centuries or perhaps decades are aout the fastest %e need1 This fast end of the
spectrum of natural clocks : tree rin!s and caron datin! : is useful for archaeolo!ical purposes0 and
for datin! specimens on the sort of timescale that covers the domestication of the do! or the
caa!e1 &t the other end of the scale0 %e need natural clocks that can time hundreds of millions0
even illions0 of years1 &nd0 praise e0 nature has provided us %ith Hust the %ide ran!e of clocks
that %e need1 What(s more0 their ran!es of sensitivity overlap %ith each other0 so %e can use them
as checks on each other1
& tree:rin! clock can e used to date a piece of %ood0 say a eam in a Tudor house0 %ith
astonishin! accuracy0 literally to the nearest year1 )ere(s ho% it %orks1 ,irst0 as most people kno%0
you can a!e a ne%ly felled tree y countin! rin!s in its trunk0 assumin! that the outermost rin!
represents the present1 Rin!s represent differential !ro%th in different seasons of the year : %inter
or summer0 dry season or %et season : and they are especially pronounced at hi!h latitudes0 %here
there is a stron! difference et%een seasons1 ,ortunately0 you don(t actually have to cut the tree
do%n in order to a!e it1 6ou can peek at its rin!s %ithout killin! it0 y orin! into the middle of a
tree and extractin! a core sample1 But Hust countin! rin!s doesn(t tell you in %hich century your
house eam %as alive0 or your +ikin! lon!ship(s mast1 #f you %ant to pin do%n the date of old0
lon!:dead %ood you need to e more sutle1 'on(t Hust count rin!s0 look at the pattern of thick and
thin rin!s1
.ust as the existence of rin!s si!nifies seasonal cycles of rich and poor !ro%th0 so some
years are etter than others0 ecause the %eather varies from year to yearA there are drou!hts that
retard !ro%th0 and umper years that accelerate itJ there are cold years and hot years0 even years of
freak El *inos or Drakatoa:type catastrophes1 Good years0 from the tree(s point of vie%0 produce
%ider rin!s than ad years1 &nd the pattern of %ide and narro% rin!s in any one re!ion0 caused y a
particular trademark seEuence of !ood years and ad years0 is sufficiently characteristic : a
fin!erprint that laels the exact years in %hich the rin!s %ere laid do%n : to e reco!niFale from
tree to tree1
'endrochronolo!ists measure rin!s on recent trees0 %here the exact date of every rin! is
kno%n y countin! ack%ards from the year in %hich the tree is kno%n to have een felled1 ,rom
these measurements0 they construct a reference collection of rin! patterns0 to %hich you can
compare the rin! patterns of an archaeolo!ical sample of %ood %hose date you %ant to kno%1 So
you mi!ht !et the reportA (This Tudor eam contains a si!nature seEuence of rin!s that matches a
seEuence from the reference collection0 %hich is kno%n to have een laid do%n in the years 2=>2 to
2=>@1 The house %as therefore uilt after &' 2=>@1(
&ll very %ell0 ut not many of today(s trees %ere alive in Tudor times0 let alone in the stone
a!e or eyond1 There are some trees : ristlecone pines0 some !iant red%oods : that live for
millennia0 ut most trees used for timer are felled %hen they are youn!er than a century or so1
)o%0 then0 do %e uild up the reference collection of rin!s for more ancient timesB ,or times so
distant that not even the oldest survivin! ristlecone pine !oes ack that farB # think you(ve already
!uessed the ans%er1 Overlaps1 & stron! rope may e 255 yards lon!0 yet no sin!le fire %ithin it
reaches more than a fraction of that total1 To use the overlap principle in dendrochronolo!y0 you
take the reference fin!erprint patterns %hose date is kno%n from modern trees1 Then you identify a
fin!erprint from the old rin!s of modern trees and seek the same fin!erprint from the youn!er rin!s
of lon!:dead trees1 Then you look at the fin!erprints from the older rin!s of those same lon!:dead
trees0 and look for the same pattern in the youn!er rin!s of even older trees1 &nd so on1 6ou can
daisychain your %ay ack0 theoretically for millions of years usin! petrified forests0 althou!h in
practice dendrochronolo!y is only used on archaeolo!ical timescales over some thousands of years1
&nd the amaFin! thin! aout dendrochronolo!y is that0 theoretically at least0 you can e accurate to
the nearest year0 even in a petrified forest 255 million years old1 6ou could literally say that this
rin! in a .urassic fossil tree %as laid do%n exactly 3=@ years later than this other rin! in another
.urassic treeK #f only there %ere enou!h petrified forests to daisychain your %ay ack continuously
from the present0 you could say that this tree is not Hust of late .urassic a!eA it %as alive in exactly
2=20>430<=@ BCK $nfortunately0 %e don(t have an unroken chain0 and dendrochronolo!y in
practice takes us ack only aout 220=55 years1 #t is nevertheless a tantaliFin! thou!ht that0 if only
%e could find enou!h petrified forests0 %e could date to the nearest year over a timespan of
hundreds of millions of years1
Ho, %en%rohronolo'y ,or&s
Tree rin!s are not Euite the only system that promises total accuracy to the nearest year1
+arves are layers of sediment laid do%n in !lacial lakes1 -ike tree rin!s0 they vary seasonally and
from year to year0 so theoretically the same principle can e used0 %ith the same de!ree of accuracy1
Coral reefs0 too0 have annual !ro%th rin!s0 Hust like trees1 ,ascinatin!ly0 these have een used to
detect the dates of ancient earthEuakes1 Tree rin!s too0 y the %ay0 tell us the dates of earthEuakes1
"ost of the other datin! systems that are availale to us0 includin! all the radioactive clocks that %e
actually use over timescales of tens of millions0 hundreds of millions or illions of years0 are
accurate only %ithin an error ran!e that is approximately proportional to the timescale concerned1
-et(s no% turn to radioactive clocks1 There are Euite a lot of them to choose from0 and0 as #
said0 they lessedly cover the !amut from centuries to thousands of millions of years1 Each one has
its o%n mar!in of error0 %hich is usually aout 2 per cent1 So if you %ant to date a rock %hich is
illions of years old0 you must e satisfied %ith an error of plus or minus tens of millions of years1
#f you %ant to date a rock hundreds of millions of years old0 you must e satisfied %ith an error of
millions1 To date a rock that is only tens of millions of years old0 you must allo% for an error of plus
or minus hundreds of thousands of years1
To understand ho% radioactive clocks %ork0 %e first need to understand %hat is meant y a
radioactive isotope1 &ll matter is made up of elements0 %hich are usually chemically comined %ith
other elements1 There are aout 255 elements0 sli!htly more if you count elements that are only ever
detected in laoratories0 sli!htly fe%er if you count only those elements that are found in nature1
Examples of elements are caron0 iron0 nitro!en0 aluminium0 ma!nesium0 fluorine0 ar!on0 chlorine0
sodium0 uranium0 lead0 oxy!en0 potassium and tin1 The atomic theory0 %hich # think everyody
accepts0 even creationists0 tells us that each element has its o%n characteristic atom0 %hich is the
smallest particle into %hich you can divide an element %ithout it ceasin! to e that element1 What
does an atom look like0 say an atom of lead0 or copper0 or caronB Well0 it certainly looks nothin!
like lead or copper or caron1 #t doesn(t look like anythin!0 ecause it is too small to form any kind
of ima!e on your retina0 even %ith an ultra:po%erful microscope1 We can use analo!ies or models
to help us visualiFe an atom1 The most famous model %as proposed y the !reat 'anish physicist
*iels Bohr1 The Bohr model0 %hich is no% rather out of date0 is a miniature solar system1 The role
of the sun is played y the nucleus0 and around it orit the electrons0 %hich play the role of planets1
&s %ith the solar system0 almost all the mass of the atom is contained in the nucleus 7(sun(80 and
almost all the volume is contained in the empty space that separates the electrons 7(planets(8 from
the nucleus1 Each electron is tiny compared %ith the nucleus0 and the space et%een them and the
nucleus is hu!e compared %ith the siFe of either1 & favourite analo!y portrays the nucleus as a fly
in the middle of a sports stadium1 The nearest nei!hourin! nucleus is another fly0 in the middle of
an adHacent stadium1 The electrons of each atom are uFFin! aout in orit around their respective
flies0 smaller than the tiniest !nats0 too small to e seen on the same scale as the flies1 When %e
look at a solid lump of iron or rock0 %e are (really( lookin! at %hat is almost entirely empty space1 #t
looks and feels solid and opaEue ecause our sensory systems and rains find it convenient to treat
it as solid and opaEue1 #t is convenient for the rain to represent a rock as solid ecause %e can(t
%alk throu!h it1 (Solid( is our %ay of experiencin! thin!s that %e can(t %alk throu!h or fall throu!h0
ecause of the electroma!netic forces et%een atoms1 (OpaEue( is the experience %e have %hen
li!ht ounces off the surface of an oHect0 and none of it !oes throu!h1
Three kinds of particle enter into the makeup of an atom0 at least as envisa!ed in the Bohr
model1 Electrons %e have already met1 The other t%o0 vastly lar!er than electrons ut still tiny
compared %ith anythin! %e can ima!ine or experience %ith our senses0 are called protons and
neutrons0 and they are found in the nucleus1 They are almost the same siFe as each other1 The
numer of protons is fixed for any !iven element and eEual to the numer of electrons1 This numer
is called the atomic numer1 #t is uniEuely characteristic of an element0 and there are no !aps in the
list of atomic numers : the famous periodic tale1L Every numer in the seEuence corresponds to
exactly one0 and only one0 element1 The element %ith 2 for its atomic numer is hydro!en0 3 is
helium0 4 lithium0 > eryllium0 = oron0 < caron0 @ nitro!en0 ; oxy!en0 and so on up to hi!h
numers like 930 %hich is the atomic numer of uranium1
Protons and electrons carry an electric char!e0 of opposite si!n : %e call one of them
positive and the other ne!ative y aritrary convention1 These char!es are important %hen elements
form chemical compounds %ith each other0 mostly mediated y electrons1 The neutrons in an atom
are ound into the nucleus to!ether %ith the protons1 $nlike protons they carry no char!e0 and they
play no role in chemical reactions1 The protons0 neutrons and electrons in any one element are
exactly the same as those in every other element1 There is no such thin! as a !old:flavoured proton
or a copper:flavoured electron or a potassium:flavoured neutron1 & proton is a proton is a proton0
and %hat makes a copper atom copper is that there are exactly 39 protons 7and exactly 39
electrons81 What %e ordinarily think of as the nature of copper is a matter of chemistry1 Chemistry
is a dance of electrons1 #t is all aout the interactions of atoms via their electrons1 Chemical onds
are easily roken and remade0 ecause only electrons are detached or exchan!ed in chemical
reactions1 The forces of attraction %ithin atomic nuclei are much harder to reak1 That(s %hy
(splittin! the atom( has such a menacin! rin! to it : ut it can happen0 in (nuclear( as opposed to
chemical reactions0 and radioactive clocks depend upon it1
Electrons have ne!li!ile mass0 so the total mass of an atom0 its (mass numer(0 is eEual to
the comined numer of protons and neutrons1 #t is usually rather more than doule the atomic
numer0 ecause there are usually a fe% more neutrons than protons in a nucleus1 $nlike the
numer of protons0 the numer of neutrons in an atom is not dia!nostic of an element1 &toms of any
!iven element can come in different versions called isotopes0 %hich have differin! numers of
neutrons0 ut al%ays the same numer of protons1 Some elements0 such as fluorine0 have only one
naturally occurrin! isotope1 The atomic numer of fluorine is 9 and its mass numer is 290 from
%hich you can deduce that it has 9 protons and 25 neutrons1 Other elements have lots of isotopes1
-ead has five commonly occurrin! isotopes1 &ll have the same numer of protons 7and electrons80
namely ;30 %hich is the atomic numer of lead0 ut the mass numers ran!e et%een 353 and 35;1
Caron has three naturally occurrin! isotopes1 Caron:23 is the common one0 %ith the same
numer of neutrons as protonsA <1 There(s also caron:240 %hich is too short:lived to other %ith0
and caron:2> %hich is rare ut not too rare to e useful for datin! relatively youn! or!anic
samples0 as %e shall see1
*o% for the next important ack!round fact1 Some isotopes are stale0 others unstale1
-ead:353 is an unstale isotopeJ lead:35>0 lead:35<0 lead:35@ and lead:35; are stale isotopes1
($nstale( means that the atoms spontaneously decay into somethin! else0 at a predictale rate0
thou!h not at predictale moments1 The predictaility of the rate of decay is the key to all
radiometric clocks1 &nother %ord for (unstale( is (radioactive(1 There are several kinds of
radioactive decay0 %hich offer possiilities for useful clocks1 ,or our purposes it isn(t important to
understand them0 ut # explain them here to sho% the ma!nificent level of detail that physicists have
achieved in %orkin! out such thin!s1 Such detail casts a sardonic li!ht on the desperate attempts of
creationists to explain a%ay the evidence of radioactive datin!0 and keep the Earth youn! like Peter
&ll these kinds of instaility involve neutrons1 #n one kind0 a neutron turns into a proton1
This means that the mass numer stays the same 7since protons and neutrons have the same mass8
ut the atomic numer !oes up y one0 so the atom ecomes a different element0 one step hi!her in
the periodic tale1 ,or example0 sodium:3> turns itself into ma!nesium:3>1 #n another kind of
radioactive decay0 exactly the reverse happens1 & proton turns into a neutron1 &!ain0 the mass
numer stays the same0 ut this time the atomic numer decreases y one0 and the atom chan!es
into the next element do%n in the periodic tale1 & third kind of radioactive decay has the same
result1 & stray neutron happens to hit a nucleus and knocks out one proton0 takin! its place1 &!ain0
there(s no chan!e in mass numerJ a!ain0 the atomic numer !oes do%n y one0 and the atom turns
into the next element do%n in the periodic tale1 There(s also a more complicated kind of decay in
%hich an atom eHects a so:called alpha particle1 &n alpha particle consists of t%o protons and t%o
neutrons stuck to!ether1 This means that the mass numer !oes do%n y four and the atomic
numer !oes do%n y t%o1 The atom chan!es to %hichever element is t%o elo% it in the periodic
tale1 &n example of alpha decay is the chan!e of the very radioactive isotope uranium:34; 7%ith
93 protons and 2>< neutrons8 to thorium:34> 7%ith 95 protons and 2>> neutrons81
*o% %e approach the nu of the %hole matter1 Every unstale or radioactive isotope decays
at its o%n characteristic rate %hich is precisely kno%n1 "oreover0 some of these rates are vastly
slo%er than others1 #n all cases the decay is exponential1 Exponential means that if you start %ith0
say0 255 !rams of a radioactive isotope0 it is not the case that a fixed amount0 say 25 !rams0 turns
into another element in a !iven time1 Rather0 a fixed proportion of %hatever is left turns into the
second element1 The favoured measure of decay rate is the (half:life(1 The half:life of a radioactive
isotope is the time taken for half of its atoms to decay1 The half:life is the same0 no matter ho%
many atoms have already decayed : that is %hat exponential decay means1 6ou %ill appreciate that0
%ith such successive halvin!s0 %e never really kno% %hen there is none left1 )o%ever0 %e can say
that after a sufficient time has elapsed : say ten half:lives : the numer of atoms that remains is so
small that0 for practical purposes0 it has all !one1 ,or example0 the half:life of caron:2> is et%een
=0555 and <0555 years1 ,or specimens older than aout =50555:<50555 years0 caron datin! is
useless0 and %e need to turn to a slo%er clock1
The half:life of ruidium:;@ is >9 illion years1 The half:life of fermium:3>> is 414
milliseconds1 Such startlin! extremes serve to illustrate the stupendous ran$e of clocks availale1
&lthou!h caron:2=(s half:life of 31> seconds is too short for settlin! evolutionary Euestions0
caron:2>(s half:life of =0@45 years is Hust ri!ht for datin! on the archaeolo!ical timescale0 and %e(ll
come to it presently1 &n isotope much used on the evolutionary timescale is potassium:>50 %ith its
half:life of 213< illion years0 and #(m !oin! to use it as my example0 to explain the %hole idea of a
radioactive clock1 #t is often called the potassium ar!on clock0 ecause ar!on:>5 7one lo%er in the
periodic tale8 is one of the elements to %hich potassium:>5 decays 7the other0 resultin! from a
different kind of radioactive decay0 is calcium:>50 one hi!her in the periodic tale81 #f you start %ith
some Euantity of potassium:>50 after 213< illion years half of the potassium:>5 %ill have decayed
to ar!on:>51 That(s %hat half:life means1 &fter another 213< illion years0 half of %hat remains 7a
Euarter of the ori!inal8 %ill have decayed0 and so on1 &fter a shorter time than 213< illion years0 a
proportionately smaller Euantity of the ori!inal potassium %ill have decayed1 So0 ima!ine that you
start %ith some Euantity of potassium:>5 in an enclosed space %ith no ar!on:>51 &fter a fe%
hundreds of millions of years have elapsed0 a scientist comes upon the same enclosed space and
measures the relative proportions of potassium:>5 and ar!on:>51 ,rom this proportion : re!ardless
of the asolute Euantities involved : kno%in! the half:life of potassium:>5(s decay and assumin!
there %as no ar!on to e!in %ith0 one can estimate the time that has elapsed since the process
started : since the clock %as (Feroed(0 in other %ords1 *otice that %e must kno% the ratio of parent
7potassium:>58 to dau!hter 7ar!on:>58 isotopes1 "oreover0 as %e sa% earlier in the chapter0 it is
necessary that our clock has the facility to e Feroed1 But %hat does it mean to speak of a
radioactive clock(s ein! (Feroed(B The process of crystalliFation !ives it meanin!1
-ike all the radioactive clocks used y !eolo!ists0 potassiumG ar!on timin! %orks only %ith
so:called i!neous rocks1 *amed after the -atin for fire0 i!neous rocks are solidified from molten
rock : under!round ma!ma in the case of !ranite0 lava from volcanoes in the case of asalt1 When
molten rock solidifies to form !ranite or asalt0 it does so in the form of crystals1 These are
normally not i!0 transparent crystals like those of EuartF0 ut crystals that are too small to look like
crystals to the naked eye1 The crystals are of various types0 and several of these0 such as some
micas0 contain potassium atoms1 &mon! these are atoms of the radioactive isotope potassium:>51
When a crystal is ne%ly formed0 at the moment %hen molten rock solidifies0 there is potassium:>5
ut no ar!on1 The clock is (Feroed( in the sense that there are no ar!on atoms in the crystal1 &s the
millions of years !o y0 the potassium:>5 slo%ly decays and0 one y one0 atoms of ar!on:>5 replace
potassium:>5 atoms in the crystal1 The accumulatin! Euantity of ar!on:>5 is a measure of the time
that has elapsed since the rock %as formed1 But0 for the reason # have Hust explained0 this Euantity is
meanin!ful only if expressed as the ratio of potassium:>5 to ar!on:>51 When the clock %as Feroed0
the ratio %as 255 per cent in favour of potassium:>51 &fter 213< illion years0 the ratio %ill e
=5:=51 &fter another 213< illion years0 half of the remainin! potassium:>5 %ill have een
converted to ar!on:>50 and so on1 #ntermediate proportions si!nify intermediate times since the
crystal clock %as Feroed1 So !eolo!ists0 y measurin! the ratio et%een potassium:>5 and ar!on:>5
in a piece of i!neous rock that they pick up today0 can tell ho% lon! a!o the rock first crystalliFed
out of its molten state1 #!neous rocks typically contain many different radioactive isotopes0 not Hust
potassium:>51 & fortunate aspect of the %ay i!neous rocks solidify is that they do so suddenly : so
that all the clocks in a !iven piece of rock are Feroed simultaneously1
Only i!neous rocks provide radioactive clocks0 ut fossils are almost never found in i!neous
rock1 ,ossils are formed in sedimentary rocks like limestone and sandstone0 %hich are not solidified
lava1 They are layers of mud or silt or sand0 !radually laid do%n on the floor of a sea or lake or
estuary1 The sand or mud ecomes compacted over the a!es and hardens as rock1 Corpses that are
trapped in the mud have a chance of fossiliFin!1 Even thou!h only a small proportion of corpses
actually do fossiliFe0 sedimentary rocks are the only rocks that contain any fossils %orth speakin!
Sedimentary rocks unfortunately cannot e dated y radioactivity1 Presumaly the
individual particles of silt or sand that !o to make sedimentary rocks contain potassium:>5 and
other radioactive isotopes0 and therefore could e said to contain radioactive clocksJ ut
unfortunately these clocks are no use to us ecause they are not properly Feroed0 or are Feroed at
different times from each other1 The particles of sand that are compacted to make sandstone may
ori!inally have een !round do%n from i!neous rocks0 ut the i!neous rocks from %hich they %ere
!round all solidified at different times1 Every !rain of sand has a clock Feroed at its o%n time0 and
that time %as proaly lon! efore the sedimentary rock formed and entomed the fossil %e are
tryin! to date1 So0 from a timekeepin! point of vie%0 sedimentary rock is a mess1 #t can(t e used1
The est %e can do : and it is a pretty !ood est : is to use the dates of i!neous rocks that are found
near sedimentary rock0 or emedded in it1
To date a fossil0 you don(t literally need to find it sand%iched et%een t%o slas of i!neous
rock0 althou!h that is a neat %ay to illustrate the principle1 The actual method used is more refined
than that1 Reco!niFaly similar layers of sedimentary rock occur all over the %orld1 -on! efore
radioactive datin! %as discovered0 these layers had een identified and !iven namesA names like
Camrian0 Ordovician0 'evonian0 .urassic0 Cretaceous0 Eocene0 Oli!ocene0 "iocene1 'evonian
sediments are reco!niFaly 'evonian0 not only in 'evon 7the county in south:%est En!land that
!ave them their name8 ut in other parts of the %orld1 They are reco!niFaly similar to each other0
and they contain similar lists of fossils1 Geolo!ists have lon! kno%n the order in %hich these
named sediments %ere laid do%n1 #t(s Hust that0 efore the advent of radioactive clocks0 %e didn(t
kno% when they %ere laid do%n1 We could arran!e them in order ecause : oviously : older
sediments tend to lie eneath youn!er sediments1 'evonian sediments0 for example0 are older than
Caroniferous 7named after the coal %hich is freEuently found in Caroniferous layers8 and %e
kno% this ecause0 in those parts of the %orld %here the t%o layers coincide0 the 'evonian layer
lies underneath the Caroniferous layer 7the exceptions to this rule occur in places %here %e can
tell0 from other evidence0 that the rocks have een tilted aslant0 or even turned upside do%n81 We
aren(t usually fortunate enou!h to find a complete run of layers0 all the %ay from Camrian at the
ottom up to Recent at the top1 But ecause the layers are so reco!niFale0 you can %ork out their
relative a!es y daisychainin! and Hi!sa%in! your %ay around the %orld1
So0 lon! efore %e kne% ho% old fossils %ere0 %e kne% the order in %hich they %ere laid
do%n0 or at least the order in %hich the named sediments %ere laid do%n1 We kne% that Camrian
fossils0 the %orld over0 %ere older than Ordovician ones0 %hich %ere older than SilurianJ then came
'evonian0 then Caroniferous0 Permian0 Triassic0 .urassic0 Cretaceous0 and so on1 &nd %ithin these
maHor named layers0 !eolo!ists also distin!uish su:re!ionsA upper .urassic0 middle .urassic0 lo%er
.urassic0 and so on1
The named strata are usually identified y the fossils they contain1 &nd %e are !oin! to use
the orderin! of the fossils as evidence for evolutionK #s that in dan!er of turnin! into a circular
ar!umentB Certainly not1 Think aout it1 Camrian fossils are a characteristic assemla!e0
unmistakaly reco!niFale as Camrian1 ,or the moment %e are usin! a characteristic assemla!e
of fossils simply as labels for Camrian rocks : indicator species : %herever %e may find them1
This0 indeed0 is %hy oil companies employ fossil experts to identify particular strata of rocks0
usually y microfossils0 tiny creatures called foraminifera0 for example0 or radiolaria1
& characteristic list of fossils is used to reco!niFe Ordovician rocks0 'evonian rocks0 and so
on1 So far0 all %e are usin! these fossil assemla!es for is to identify %hether a sla of rock is0 say0
Permian or Silurian1 *o% %e move on to use the order in %hich the named strata %ere laid do%n0
helped y daisychainin! around the %orld0 as evidence of %hich strata are older or youn!er than
%hich1 )avin! estalished these t%o sets of information0 %e can then look at the fossils in
successively youn!er strata0 to see %hether they constitute a sensile evolutionary seEuence %hen
compared %ith each other in seEuence1 'o they pro!ress in a sensile directionB 'o certain kinds of
fossils0 for example mammals0 appear only after a !iven date0 never eforeB The ans%er to all such
Euestions is yes1 &l%ays yes1 *o exceptions1 That is po%erful evidence for evolution0 for it %as
never a necessary fact0 never somethin! that had to follo% from our method of identifyin! strata
and our method of otainin! a temporal seEuence1
#t is a fact that literally nothin! that you could remotely call a mammal has ever een found
in 'evonian rock or in any older stratum1 They are not Hust statistically rarer in 'evonian than in
later rocks1 They literally never occur in rocks older than a certain date1 But this didn(t have to e
so1 #t could have een the case that0 as %e du! do%n lo%er and lo%er from the 'evonian0 throu!h
the Silurian and then even older0 throu!h the Ordovician0 %e suddenly found that the Camrian era
: older than any of them : teemed %ith mammals1 That is in fact not %hat %e find0 ut the
possiility demonstrates that you can(t accuse the ar!ument of ein! circularA at any moment
someody mi!ht di! up a mammal in Camrian rocks0 and the theory of evolution %ould e
instantly lo%n apart if they did1 Evolution0 in other %ords0 is a falsifiale0 and therefore scientific0
theory1 # shall return to this point in Chapter <1
Creationist attempts to explain such findin!s often achieve hi!h comedy1 *oah(s flood0 %e
are told0 is the key to understandin! the order in %hich %e find fossils of the maHor animal !roups1
)ere(s a direct Euotation from a priFe%innin! creationist %esite1
,ossil seEuence in !eolo!ical strata sho%sA
7i8 #*+ERTEBR&TES 7slo% movin! marine animals8 %ould perish first follo%ed y the
more moile fishes %ho %ould e over%helmed y the flood silt
7ii8 &"P)#B#& 7close to the sea8 %ould perish next as the %aters rose1
7iii8 REPT#-ES 7slo% movin! land animals8 next to die1
7iv8 "&""&-S could flee from risin! %ater0 the lar!er0 faster ones survivin! the lon!est1
7v8 "&* %ould exercise most in!enuity : clin!in! to lo!s0 etc1 to escape the flood1
This seEuence is a perfectly satisfactory explanation of the order in %hich the various fossils
are found in the strata1 #t is *OT the order in %hich they evolved ut the order in %hich they %ere
inundated at the time of *oah(s flood1
Puite apart from all the other reasons to oHect to this remarkale explanation0 there could
only ever e a statistical tendency for mammals0 for example0 to e on aera$e etter at escapin!
the risin! %aters than reptiles1 #nstead0 as %e should expect on the evolution theory0 there literally
are no mammals in the lo%er strata of the !eolo!ical record1 The (head for the hills( theory %ould e
on more solid !round if there %ere a statistical tailin! off of mammals as you move do%n throu!h
the rocks1 There are literally no triloites aove Permian strata0 literally no dinosaurs 7except irds8
aove Cretaceous strata1 Once a!ain0 the (head for the hills( theory %ould predict a statistical tailin!
Back to datin!0 and radioactive clocks1 Because the relative orderin! of the named
sedimentary strata is %ell kno%n0 and the same order is found all over the %orld0 %e can use
i!neous rocks that overlie or underlie sedimentary strata0 or are emedded in them0 to date those
named sedimentary strata0 and hence the fossils %ithin them1 By a refinement of the method0 %e can
date fossils that lie near the top of0 say0 the Caroniferous or the Cretaceous0 as more recent than
fossils that lie sli!htly lo%er in the same stratum1 We don(t need to find an i!neous rock in the
vicinity of any particular fossil %e %ant to date1 We can tell that our fossil is0 say0 late 'evonian0
from its position in a 'evonian stratum1 &nd %e kno%0 from the radioactive datin! of i!neous rocks
found in association %ith 'evonian strata all around the %orld0 that the 'evonian ended aout 4<5
million years a!o1
Ra%ioati.e lo&s
The potassium ar!on clock is only one of many clocks that are availale to !eolo!ists0 all
usin! the same principle on their different timescales1 &ove is a tale of clocks0 ran!in! from slo%
to fast1 *otice0 yet a!ain0 the astonishin! ran!e of half:lives0 from >9 illion years at the slo% end to
less than <0555 years at the fast end1 The faster clocks0 such as caron:2>0 %ork in a some%hat
different %ay1 This is ecause the (Feroin!( of these hi!her:speed clocks is necessarily different1 ,or
isotopes %ith a short half:life0 all the atoms that %ere present %hen the Earth %as ori!inally formed
have lon! since disappeared1 Before # turn to ho% caron datin! %orks0 it is %orth pausin! to
consider another piece of evidence in favour of an old Earth0 a planet %hose a!e is measured in
illions of years1
&mon! all the elements that occur on Earth are 2=5 stale isotopes and 2=; unstale ones0
makin! 45; in all1 Of the 2=; unstale ones0 232 are either extinct or exist only ecause they are
constantly rene%ed0 like caron:2> 7as %e shall see81 *o%0 if %e consider the 4@ that have not !one
extinct0 %e notice somethin! si!nificant1 Every sin!le one of them has a half:life !reater than @55
million years1 &nd if %e look at the 232 that have !one extinct0 every sin!le one of them has a half:
life less than 355 million years1 'on(t e misled0 y the %ay1 Rememer %e are talkin! half0life
here0 not lifeK Think of the fate of an isotope %ith a half:life of 255 million years1 #sotopes %hose
half:life is less than a tenth or so of the a!e of the Earth are0 for practical purposes0 extinct0 and
don(t exist except under special circumstances1 With exceptions that are there for a special reason
that %e understand0 the only isotopes that %e find on Earth are those that have a half:life lon!
enou!h to have survived on a very old planet1 Caron:2> is one of these exceptions0 and it is
exceptional for an interestin! reason0 namely that it is ein! continuously replenished1 Caron:2>(s
role as a clock therefore needs to e understood in a different %ay from that of lon!er:lived
isotopes1 #n particular0 %hat does it mean to 2ero the clockB
Of all the elements0 caron is the one that seems most indispensale to life : the one %ithout
%hich life on any planet is hardest to envisa!e1 This is ecause of caron(s remarkale capacity for
formin! chains and rin!s and other complex molecular architectures1 #t enters the food %e via
photosynthesis0 %hich is the process %herey !reen plants take in caron dioxide molecules from
the atmosphere and use ener!y from sunli!ht to comine the caron atoms %ith %ater to make
su!ars1 &ll the caron in ourselves and in all other livin! creatures comes ultimately0 via plants0
from caron dioxide in the atmosphere1 &nd it is continually ein! recycled ack to the atmosphereA
%hen %e reathe out0 %hen %e excrete0 and %hen %e die1
"ost of the caron in the atmosphere(s caron dioxide is caron:230 %hich is not
radioactive1 )o%ever0 aout one atom in a trillion is caron:2>0 %hich is radioactive1 #t decays
rather rapidly0 %ith a half:life of =0@45 years0 as %e have seen0 to nitro!en:2>1 Plant iochemistry is
lind to the difference et%een these t%o carons1 To a plant0 caron is caron is caron1 So plants
take in caron:2> alon!side caron:230 and incorporate the t%o kinds of caron atom in su!ars0 in
the same proportion as they exist in the atmosphere1 The caron that is incorporated from the
atmosphere 7complete %ith the same proportion of caron:2> atoms8 is rapidly 7compared to the
half:life of caron:2>8 spread throu!h the food chain0 as plants are eaten y herivores0 herivores
y carnivores and so on1 &ll livin! creatures0 %hether plants or animals0 have approximately the
same ratio of caron:23 to caron:2>0 %hich is the same ratio as you(ll find in the atmosphere1
So0 %hen is the clock FeroedB &t the moment %hen a livin! creature0 %hether animal or
plant0 dies1 &t that moment0 it is severed from the food chain0 and detached from the inflo% of fresh
caron:2>0 via plants0 from the atmosphere1 &s the centuries !o y0 the caron:2> in the corpse0 or
lump of %ood0 or piece of cloth0 or %hatever it is0 steadily decays to nitro!en:2>1 The ratio of
caron:2> to caron:23 in the specimen therefore !radually drops further and further elo% the
standard ratio that livin! creatures share %ith the atmosphere1 Eventually it %ill e all caron:23 :
or0 more strictly0 the caron:2> content %ill ecome too small to measure1 &nd the ratio of
caron:23 to caron:2> can e used to calculate the time that has elapsed since the death of the
creature cut it off from the food chain and its interchan!e %ith the atmosphere1
That(s all very %ell0 ut it only %orks ecause there is a continuously replenished supply of
caron:2> in the atmosphere1 Without that0 the caron:2> %ith its short half:life %ould lon! since
have disappeared from the Earth0 alon! %ith all other naturally occurrin! isotopes %ith short half:
lives1 Caron:2> is special ecause it is continually ein! made y cosmic rays omardin!
nitro!en atoms in the upper atmosphere1 *itro!en is the commonest !as in the atmosphere and its
mass numer is 2>0 the same as caron:2>(s1 The difference is that caron:2> has < protons and ;
neutrons0 %hile nitro!en:2> has @ protons and @ neutrons 7neutrons0 rememer0 have near:enou!h
the same mass as protons81 Cosmic ray particles are capale of hittin! a proton in a nitro!en nucleus
and convertin! it to a neutron1 When this happens0 the atom ecomes caron:2>0 caron ein! one
lo%er than nitro!en in the periodic tale1 The rate at %hich this conversion occurs is approximately
constant from century to century0 %hich is %hy caron datin! %orks1 &ctually the rate is not exactly
constant0 and ideally %e need to compensate for this1 ,ortunately %e have an accurate caliration of
the fluctuatin! supply of caron:2> in the atmosphere and can take this into account to refine our
datin! calculations1 Rememer that0 over rou!hly the same a!e ran!e as is covered y caron
datin!0 %e have an alternative method of datin! %ood : dendrochronolo!y : %hich is completely
accurate to the nearest year1 By lookin! at the caron:dated a!es of %ood samples %hose a!e is
independently kno%n from tree:rin! datin!0 %e can calirate the fluctuatin! errors in caron:datin!1
Then %e can use these caliration measurements %hen %e !o ack to or!anic samples for %hich %e
don(t have tree:rin! data 7the maHority81
Caron datin! is a comparatively recent invention0 !oin! ack only to the 29>5s1 #n its early
years0 sustantial Euantities of or!anic material %ere needed for the datin! procedure1 Then0 in the
29@5s0 a techniEue called mass spectrometry %as adapted to caron datin!0 %ith the result that only
a tiny Euantity of or!anic material is no% needed1 This has revolutioniFed archaeolo!ical datin!1
The most celerated example is the Shroud of Turin1 Since this notorious piece of cloth seems
mysteriously to have imprinted on it the ima!e of a earded0 crucified man0 many people hoped it
mi!ht hail from the time of .esus1 #t first turns up in the historical record in the mid:fourteenth
century in ,rance0 and noody kno%s %here it %as efore that1 #t has een housed in Turin since
2=@;0 under the custody of the +atican since 29;41 When mass spectrometry made it possile to
date a tiny sample of the shroud0 rather than the sustantial s%athes that %ould have een needed
efore0 the +atican allo%ed a small strip to e cut off1 The strip %as divided into three parts and
sent to three leadin! laoratories specialiFin! in caron datin!0 in Oxford0 &riFona and Iurich1
Workin! under conditions of scrupulous independence : not comparin! notes : the three
laoratories reported their verdicts on the date %hen the flax from %hich the cloth had een %oven
died1 Oxford said ad 23550 &riFona 245> and Iurich 23@>1 These dates are all : %ithin normal
mar!ins of error : compatile %ith each other and %ith the date in the 24=5s at %hich the shroud is
first mentioned in history1 The datin! of the shroud remains controversial0 ut not for reasons that
cast dout on the caron:datin! techniEue itself1 ,or example0 the caron in the shroud mi!ht have
een contaminated y a fire0 %hich is kno%n to have occurred in 2=431 # %on(t pursue the matter
further0 ecause the shroud is of historical0 not evolutionary0 interest1 #t is a nice example0 ho%ever0
to illustrate the method0 and the fact that0 unlike dendrochronolo!y0 it is not accurate to the nearest
year0 only to the nearest century or so1
# have repeatedly emphasiFed that there are lots of different clocks that the modern
evolutionary detective can use0 and also that they %ork est on different0 ut overlappin!
timescales1 Radioactive clocks can e used to !ive independent estimates of the a!e of one piece of
rock0 earin! in mind that all the clocks %ere Feroed simultaneously %hen this very same piece of
rock solidified1 When such comparisons have een made0 the different clocks a!ree %ith each other
: %ithin the expected mar!ins of error1 This !ives !reat confidence in the correctness of the clocks1
Thus mutually calirated and verified on kno%n rocks0 these clocks can e carried %ith confidence
to interestin! datin! prolems0 such as the a!e of the Earth itself1 The currently a!reed a!e of >1<
illion years is the estimate upon %hich several different clocks conver!e1 Such a!reement is not
surprisin!0 ut unfortunately %e need to emphasiFe it ecause0 astonishin!ly0 as # pointed out in the
#ntroduction 7and have documented in the &ppendix80 some >5 per cent of the &merican population0
and a some%hat smaller percenta!e of the British population0 claim to elieve that the a!e of the
Earth0 far from ein! measured in illions of years0 is less than 250555 years1 -amentaly0
especially in &merica and over much of the #slamic %orld0 some of these history:deniers %ield
po%er over schools and their syllauses1
*o%0 a history:denier could claim0 say0 that there is somethin! %ron! %ith the potassium
ar!on clock1 What if the present very slo% rate of decay of potassium:>5 has only een in operation
since *oah(s floodB #f0 efore that0 the half:life of potassium:>5 %as radically different0 only a fe%
centuries0 say0 rather than 213< illion yearsB The special pleadin! in such claims is !larin!1 Why on
Earth should the la%s of physics chan!e0 Hust like that0 so massively and so convenientlyB &nd it
!lares even more %hen you have to make mutually adHusted special pleadin! claims for each one of
the clocks separately1 &t present0 the applicale isotopes all a!ree %ith each other in placin! the
ori!in of the Earth at et%een four and five illion years a!o1 &nd they do so on the assumption that
their half:lives have al%ays een the same as %e can measure today : as the kno%n la%s of physics0
indeed0 stron!ly su!!est they should1 The history:deniers %ould have to fiddle the half:lives of all
the isotopes in their separate proportions0 so that they all end up a!reein! that the Earth e!an <0555
years a!o1 *o% that(s %hat # call special pleadin!K &nd # haven(t even mentioned various other
datin! methods %hich also produce the same result0 for example (fission track datin!(1 Bear in mind
the hu!e differences in timescales of the different clocks0 and think of the amount of contrived and
complicated fiddlin! %ith the la%s of physics that %ould e needed in order to make all the clocks
a!ree %ith each other0 across the orders of ma!nitude0 that the Earth is <0555 years old and not >1<
illionK Given that the sole motive for such fiddlin! is the desire to uphold the ori!in myth of a
particular set of BronFe &!e desert triesmen0 it is surprisin!0 to say the least0 that anyone is fooled
y it1
There is one more type of evolutionary clock0 the molecular clock0 ut # shall postpone
discussin! it until Chapter 250 after introducin! some other ideas of molecular !enetics1
L # am a sundial0 and # make a otch
Of %hat is done far etter y a %atch
)ilaire Belloc
L Which is presumaly ased on the evolutionary happenstance of our possessin! ten
fin!ers1 ,red )oyle has in!eniously speculated that0 if %e had een orn %ith ei!ht di!its and
therefore ecome accustomed to octal arithmetic instead of decimal0 %e mi!ht have invented inary
arithmetic and hence electronic computers a century earlier than %e did 7since ; is a po%er of 381
L &las0 the popular le!end that it came to 'mitri "endeleev in a dream may e false1
# )&+E used the metaphor of a detective0 comin! on the scene of a crime after it is all over
and reconstructin! from the survivin! clues %hat must have happened1 But perhaps # %as too ready
to concede the impossiility of vie%in! evolution as an eye %itness1 &lthou!h the vast maHority of
evolutionary chan!e took place efore any human ein! %as orn0 some examples are so fast that
%e can see evolution happenin! %ith our o%n eyes durin! one human lifetime1
There(s a plausile indication that this may have happened even %ith elephants0 %hich
'ar%in himself picked out as one of the slo%est:reproducin! animals0 %ith one of the lon!est
!eneration turnovers1 One of the main causes of mortality amon! &frican elephants is humans %ith
!uns huntin! ivory0 either for trophies or to sell for carvin!1 *aturally the hunters tend to pick on
the individuals %ith the lar!est tusks1 This means that0 at least in theory0 smaller:tusked individuals
%ill e at a selective advanta!e1 &s ever %ith evolution0 there %ill e conflictin! selection
pressures0 and %hat %e see evolvin! %ill e a compromise1 -ar!er tuskers doutless have an
advanta!e %hen it comes to competition %ith other elephants0 and this %ill e alanced a!ainst their
disadvanta!e %hen they encounter men %ith !uns1 &ny increase in huntin! activity0 %hether in the
form of ille!al poachin! or le!al huntin!0 %ill tend to shift the alance of advanta!e to%ards smaller
tusks1 &ll other thin!s ein! eEual0 %e mi!ht expect an evolutionary trend to%ards smaller tusks as
a result of human huntin!0 ut %e(d proaly expect it to take millennia to e detectale1 We %ould
not expect to see it %ithin one human lifetime1 *o% let(s look at some fi!ures1
T(s& ,ei'ht in 0'an%an ele#hants
The !raph aove sho%s data from the $!anda Game 'epartment0 pulished in 29<31
Referrin! only to elephants le!ally shot y licensed hunters0 it sho%s mean tusk %ei!ht in pounds
7that dates it8 from year to year et%een 293= and 29=; 7durin! %hich time $!anda %as a British
protectorate81 The dots are annual fi!ures1 The line throu!h the dots is dra%n not y eye ut y a
statistical techniEue called linear re!ression1L 6ou can see that there is a decreasin! trend over the
thirty:three years1 &nd the trend is hi!hly statistically si!nificant0 %hich means that it is almost
certainly a real trend0 not a random chance effect1
The fact that there is a statistically si!nificant trend to%ards shrinkin! tusks doesn(t
necessarily mean it is an evolutionary trend1 #f you %ere to plot a !raph of mean hei!ht of 35:year:
old men0 from year to year durin! the t%entieth century0 you(d see in many countries a si!nificant
trend to%ards !ettin! taller1 This is normally reckoned to e not an evolutionary trend0 ut rather an
effect of improved nutrition1 *evertheless0 in the case of the elephants %e have !ood reason to
suspect the existence of stron! selection a!ainst lar!e tusks1 Reflect that0 althou!h the !raph refers
to tusks otained from licensed kills0 the selection pressure that produced the trend could %ell have
come mostly from poachin!1 We must seriously entertain the possiility that it is a true evolutionary
trend0 in %hich case it is a remarkaly rapid one1 We must e cautious efore concludin! too much1
#t could e that %e are oservin! stron! natural selection0 %hich is hi!hly likely to result in chan!es
in !ene freEuencies in the population0 ut such !enetic effects have not so far een demonstrated1 #t
could e that the difference et%een lar!e:tusked and small:tusked elephants is a non:!enetic
difference1 *evertheless0 # am inclined to take seriously the possiility that it is a true evolutionary
"ore to the point0 my collea!ue 'r #ain 'ou!las:)amilton0 %ho is the %orld authority on
%ild &frican elephant populations0 takes it seriously and elieves0 surely ri!htly0 that it needs
lookin! into more closely1 )e suspects that the trend started lon! efore 293= and has continued
after 29=;1 )e has reason to think that the same cause0 operatin! in the past0 underlies the
tusklessness of many local populations of &sian elephants1 We seem to have here a prima facie case
of rapid evolution takin! place efore our very eyes0 one that %ould repay further research1
-et me turn0 no%0 to another case0 one in %hich there is some intri!uin! recent researchA a
study of liFards on &driatic islands1
T)E -#I&R'S O, PO' "RC&R$
There are t%o small islets off the Croatian coast called Pod Dopiste and Pod "rcaru1 #n
29@2 a population of common "editerranean liFards0 Podarcis sic"la0 %hich mainly eat insects0
%as present on Pod Dopiste ut there %ere none on Pod "rcaru1 #n that year experimenters
transported five pairs of Podarcis sic"la from Pod Dopiste and released them on Pod "rcaru1 Then0
in 355;0 another !roup of mainly Bel!ian scientists0 associated %ith &nthony )errel0 visited the
islands to see %hat had happened1 They found a flourishin! population of liFards on Pod "rcaru0
%hich '*& analysis confirmed %ere indeed Podarcis sic"la1 These are presumed to have
descended from the ori!inal five pairs that %ere transported1 )errel and his collea!ues made
oservations on the descendants of the transported liFards0 and compared them %ith liFards livin!
on the ori!inal ancestral island1 There %ere marked differences1 The scientists made the proaly
Hustified assumption that the liFards on the ancestral island0 Pod Dopiste0 %ere unchan!ed
representatives of the ancestral liFards of thirty:six years efore1 #n other %ords0 they presumed they
%ere comparin! the evolved liFards of Pod "rcaru %ith their unevolved (ancestors( 7meanin! their
contemporaries ut of ancestral type8 on Pod Dopiste1 Even if this presumption is %ron! : even if0
for example0 the liFards of Pod Dopiste have een evolvin! Hust as fast as the liFards of Pod "rcaru
: %e are still oservin! evolutionary diver!ence in nature0 over a timescale of decadesA the sort of
timescale that humans can oserve %ithin one lifetime1
&nd %hat %ere the differences et%een the t%o island populations0 differences that had
taken a mere thirty:seven years or so to evolveBL Well0 the Pod "rcaru liFards : the (evolved(
population : had si!nificantly lar!er heads than the (ori!inal( Pod Dopiste populationA lon!er0 %ider
and taller heads1 This translates into a markedly !reater ite force1 & chan!e of this kind typically
!oes %ith a shift to a more ve!etarian diet and0 sure enou!h0 the liFards of Pod "rcaru eat
si!nificantly more plant material than the (ancestral( type on Pod Dopiste1 ,rom the almost
exclusive diet of insects 7arthropods0 in the terms of the !raph opposite8 still enHoyed y the modern
Pod Dopiste population0 the liFards on Pod "rcaru had shifted to a lar!ely ve!etarian diet0
especially in summer1
Why %ould an animal need a stron!er ite %hen shiftin! to a ve!etarian dietB Because plant0
ut not animal0 cell %alls are stiffened y cellulose1 )erivorous mammals like horses0 cattle and
elephants have !reat millstone:like teeth for !rindin! cellulose0 Euite different from the shearin!
teeth of carnivores and the needly teeth of insectivores1 &nd they have massive Ha% muscles0 and
correspondin!ly roust skulls for the muscle attachments 7think of the stout midline crest alon! the
top of a !orilla(s skull81 L +e!etarians also have characteristic peculiarities of the !ut1 &nimals
!enerally can(t di!est cellulose %ithout the aid of acteria or other micro:or!anisms0 and many
verterates set aside a lind alley in the !ut called the caecum0 %hich houses such acteria and acts
as a fermentation chamer 7our appendix is a vesti!e of the lar!er caecum in our more ve!etarian
ancestors81 The caecum0 and other parts of the !ut0 can ecome Euite elaorate in specialist
herivores1 Carnivores usually have simpler !uts than herivores0 and smaller too1 &mon! the
complications that ecome inserted in herivore !uts are thin!s called caecal valves1 +alves are
incomplete partitions0 sometimes muscular0 %hich can serve to re!ulate or slo% do%n the flo% of
material throu!h the !ut0 or simply increase the surface area of the interior of the caecum1 The
picture on the left sho%s the caecum cut open in a related species of liFard %hich eats a lot of plant
material1 The valve is indicated y the arro%1 *o%0 the fascinatin! thin! is that0 althou!h caecal
valves don(t normally occur in Podarcis sic"la and are rare in the family to %hich it elon!s0 those
valves have actually started to evolve in the population of P4 sic"la on Pod "rcaru0 the population
that has0 for only the past thirty:seven years0 een evolvin! to%ards herivory1 The investi!ators
discovered other evolutionary chan!es in the liFards of Pod "rcaru1 The population density
increased0 and the liFards ceased to defend territories in the %ay that the (ancestral( population on
Pod Dopiste did1 # should repeat that the only thin! that is really exceptional aout this %hole story0
and the reason # am tellin! it here0 is that it all happened so extremely rapidly0 in a matter of a fe%
decadesA evolution efore our very eyes1
S(""er %iet o$ li-ar%s on t,o A%riati islan%s
Caeal .al.e
,ORT6:,#+E T)O$S&*' GE*ER&T#O*S O, E+O-$T#O* #* T)E -&B
The avera!e !eneration turnover of those liFards is aout t%o years0 so the evolutionary
chan!e oserved on Pod "rcaru represents only aout ei!hteen or nineteen !enerations1 .ust think
%hat you mi!ht see in three or four decades if you follo%ed the evolution of acteria0 %hose
!enerations are measured in hours or even minutes0 rather than yearsK Bacteria offer another
priceless !ift to the evolutionist1 #n some cases you can freeFe them for an indefinite len!th of time
and then rin! them ack to life a!ain0 %hereupon they resume reproduction as if nothin! had
happened1 This means that experimenters can lay do%n their o%n (livin! fossil record(0 a snapshot of
the exact point the evolutionary process had reached at any desired time1 #ma!ine if %e could rin!
-ucy0 the ma!nificent pre:human fossil discovered y 'on .ohanson0 ack to life from a deep:
freeFe and set her kind evolvin! ane%K &ll this has een achieved %ith the acterium Escherichia
coli0 in a spectacular lon!:term experiment y the acteriolo!ist Richard -enski and his collea!ues
at "ichi!an State $niversity1 Scientific research no%adays is often a team effort1 #n %hat follo%s0 #
may sometimes use the name (-enski( for revity0 ut you should read it as (-enski and the
collea!ues and students in his la(1 &s %e shall see0 the -enski experiments are distressin! to
creationists0 and for a very !ood reason1 They are a eautiful demonstration of evolution in action0
somethin! it is hard to lau!h off even %hen your motivation to do so is very stron!1 &nd the
motivation for dyed:in:the:%ool creationists is very stron! indeed1 #(ll return to this at the end of the
E4 coli is a common acterium1 +ery common1 There are aout a hundred illion illion of
them around the %orld at any one time0 of %hich aout a illion0 y -enski(s calculation0 are in your
lar!e intestine at this very moment1 "ost of them are harmless or even eneficial0 ut nasty strains
occasionally hit the headlines1 Such periodic evolutionary innovation is not surprisin! if you do the
sums0 even thou!h mutations are rare events1 #f %e assume that the proaility of a !ene mutatin!
durin! any one act of acterial reproduction is as lo% as one in a illion0 the numers of acteria are
so colossal that Hust aout every !ene in the !enome %ill have mutated some%here in the %orld0
every day1 &s Richard -enski says0 (That(s a lot of opportunity for evolution1(
-enski and his collea!ues exploited that opportunity0 in a controlled %ay0 in the la1 Their
%ork is extremely thorou!h and careful in every detail1 The details really contriute to the impact of
the evidence for evolution that these experiments provide0 and # am therefore not !oin! to stint in
explainin! them1 This means that the next fe% pa!es are inevitaly some%hat intricate : not
difficult0 Hust intricately detailed1 #t %ould proaly e est not to read this section of the ook %hen
tired0 at the end of a lon! day1 What makes it easier to follo% is that every detail makes senseA none
of it leaves us scratchin! our heads and %onderin! %hat that %as all aout1 So0 please come %ith
me0 step y step0 throu!h this splendidly constructed and ele!antly executed set of experiments1
These acteria reproduce asexually : y simple cell division : so it is easy to clone up a hu!e
population of !enetically identical individuals in a short time1 #n 29;;0 -enski took one such
population and infected t%elve identical flasks0 all of %hich contained the same nutrient roth0
includin! !lucose as the vital food source1 The t%elve flasks0 each %ith its foundin! population of
acteria0 %ere then placed in a (shakin! incuator( %here they %ere kept nice and %arm0 and shaken
to keep the acteria %ell distriuted throu!hout the liEuid1 These t%elve flasks founded t%elve lines
of evolution that %ere destined to e kept separate from one another for t%o decades and countin!A
sort of like the t%elve tries of #srael0 except that in the case of the tries of #srael there %as no la%
a!ainst their mixin!1
The t%elve tries of acteria %ere not kept in the same t%elve flasks for all that time1 On the
contrary0 each trie had a ne% flask every day1 #ma!ine t%elve lines of flasks0 stretchin! a%ay into
the distance0 each line more than @0555 flasks lon!K Every day0 for each of the t%elve tries0 a ne%
vir!in flask %as infected %ith liEuid from the previous day(s flask1 & small sample0 exactly one:
hundredth of the volume of the old flask0 %as dra%n out and sEuirted into the ne% flask0 %hich
contained a fresh supply of !lucose:rich roth1 The population of acteria in the flask then started to
skyrocketJ ut it al%ays levelled off y the next day as the supply of food !ave out and starvation
set in1 #n other %ords0 the population in every flask multiplied itself hu!ely0 then reached a plateau0
at %hich point a ne% infective sample %as dra%n and the cycle rene%ed the next day1 Thousands of
times throu!h their hi!h:speed eEuivalent of !eolo!ical time0 therefore0 these acteria %ent throu!h
the same daily repeated cycles of onanFa expansion0 follo%ed y starvation0 from %hich a lucky
hundredth %ere rescued and carried0 in a !lass *oah(s &rk0 to a fresh : ut a!ain temporary :
!lucose onanFaA perfect perfect perfect conditions for evolution0 and0 %hat is more0 the experiment
%as done in t%elve separate lines in parallel1
-enski and his team have continued this daily routine for more than t%enty years so far1
This means aout @0555 (flask !enerations( and >=0555 acterial !enerations : avera!in! et%een six
and seven acterial !enerations per day1 To put that into perspective0 if %e %ere to !o ack >=0555
human !enerations0 that %ould e aout a million years0 ack to the time of )omo erect"s0 %hich is
not very lon! a!o1 So0 %hatever evolutionary chan!e -enski may have clocked up in the eEuivalent
of a million years of acterial !enerations0 think ho% much more evolution mi!ht happen in0 say0
255 million years of mammal evolution1 &nd even 255 million years is comparatively recent0 y
!eolo!ical standards1
#n addition to the main evolution experiment0 the -enski !roup used the acteria for various
illuminatin! spin:off experiments0 for example replacin! !lucose %ith another su!ar0 maltose0 after
30555 !enerations0 ut # shall concentrate on the central experiment0 %hich used !lucose throu!hout1
They sampled the t%elve tries at intervals throu!hout the t%enty years0 to see ho% evolution %as
pro!ressin!1 They also froFe samples of each of the tries as a source of resuscitatale (fossils(
representin! strate!ic points alon! the evolutionary %ay1 #t is hard to exa!!erate ho% rilliantly
conceived this series of experiments is1
)ere(s a little example of the excellent for%ard plannin!1 6ou rememer # said that the
t%elve foundin! flasks %ere all seeded from the same clone and therefore started out !enetically
identical1 But that %asn(t Euite true : for an interestin! and cunnin! reason1 The -enski la had
earlier exploited a !ene called ara %hich comes in t%o forms0 &raO and &raMBN1 6ou can(t tell the
difference until you take a sample of the acteria and (plate them out( on an a!ar plate that contains
a nutritious roth plus the su!ar arainose and a chemical dye called tetraFolium1 (Platin! out( is one
of the thin!s acteriolo!ists do1 #t means puttin! a drop of liEuid0 containin! acteria0 on a plate
covered %ith a thin sheet of a!ar !el and then incuatin! the plate1 Colonies of acteria !ro% out as
expandin! circles : miniature fairy rin!s L : from the drops0 feedin! on the nutrients mixed in %ith
the a!ar1 #f the mixture contains arainose and the indicator dye0 the difference et%een &raO and
&raMBN is revealed0 as if y heatin! invisile inkA they sho% up as %hite and red colonies0
respectively1 The -enski team find this colour distinction useful for laellin! purposes0 as %e shall
see0 and they anticipated this usefulness y settin! up six of their t%elve tries as &raO and the
other six as &raMBN1 .ust to !ive one example of ho% they exploited the colour codin! of the
acteria0 they used it as a check on their o%n laoratory procedures1 When performin! their daily
ritual of infectin! ne% flasks0 they took care to handle &raO and &raMBN flasks alternately1 That %ay0
if they ever made a mistake : splashed a transfer pipette %ith liEuid or somethin! like that : it %ould
sho% up %hen they later suHected samples to the redG%hite test1 #n!eniousB 6es1 &nd scrupulous1
Really !ood scientists have to e oth1
But for!et aout &raO and &raMBN for the moment1 #n all other respects0 the foundin!
populations of the t%elve tries started out identical1 *o other differences et%een &raMBN and &raO
have een detected0 so they really could e treated as convenient colour markers0 as ornitholo!ists
put colour rin!s on irds( le!s1
Ri!ht0 then1 We have our t%elve tries0 marchin! throu!h their o%n hi!hly speeded:up
eEuivalent of !eolo!ical time0 in parallel0 under the same conditions of repeated oom and ust1 The
interestin! Euestion %as0 %ould they stay the same as their ancestorsB Or %ould they evolveB &nd if
they evolved0 %ould all t%elve tries evolve in the same %ay0 or %ould they diver!e from one
The roth0 as # have said0 contained !lucose1 #t %as not the only food there0 ut it %as the
limitin! resource1 This means that runnin! out of !lucose %as the key factor that caused the
population siFe0 in every flask every day0 to stop climin! and reach a plateau1 To put it another
%ay0 if the experimenters had put more !lucose in the daily flasks0 the population plateau at the end
of the day %ould have een hi!her1 Or0 if they had added a second dollop of !lucose after the
plateau %as reached0 they %ould have %itnessed a second spurt of population !ro%th0 up to a ne%
#n these conditions0 the 'ar%inian expectation %as that0 if any mutation arose that assisted
an individual acterium to exploit !lucose more efficiently0 natural selection %ould favour it0 and it
%ould spread throu!h the flask as mutant individuals out:reproduced non:mutant individuals1 #ts
type %ould then disproportionately infect the next flask in the linea!e and0 as flask took over from
flask0 pretty soon the mutant %ould have a monopoly of its trie1 Well0 this is exactly %hat
happened in all t%elve tries1 &s the (flask !enerations( %ent y0 all t%elve lines improved over
their ancestorsA !ot etter at exploitin! !lucose as a food source1 But0 fascinatin!ly0 they !ot etter
in different %ays : that is0 different tries developed different sets of mutations1
)o% did the scientists kno% thisB They could tell y samplin! the linea!es as they evolved0
and comparin! the (fitness( of each sample a!ainst (fossils( sampled from the ori!inal foundin!
population1 Rememer that (fossils( are froFen samples of acteria %hich0 %hen unfroFen0 carry on
livin! and reproducin! normally1 &nd ho% did -enski and his collea!ues make this comparison of
(fitness(B )o% did they compare (modern( acteria %ith their (fossil( ancestorsB With !reat in!enuity1
They took a sample of the putatively evolved population and put it in a vir!in flask1 &nd they put a
same:siFed sample of the unfroFen ancestral population in the same flask1 *eedless to say0 these
experimentally mixed flasks %ere from then on entirely removed from contact %ith the continuin!
linea!es of the t%elve tries in the lon!:term evolution experiment1 This side experiment %as done
%ith samples that played no further part in the main experiment1
So0 %e have a ne% experimental flask containin! t%o competin! strains0 (modern( and (livin!
fossil(0 and %e %ant to kno% %hich of the t%o strains %ill out:populate the other1 But they are all
mixed up0 so ho% do you tellB )o% do you distin!uish the t%o strains0 %hen they are mixed
to!ether in the (competition flask(B # told you it %as in!enious1 6ou rememer the colour codin!0
%ith the (reds( 7&raMBN8 and the (%hites( 7&raO8B *o%0 if you %anted to compare the fitness of0 say0
Trie = %ith the ancestral fossil population0 %hat %ould you doB -et(s suppose that Trie = %as
&raO1 Well then0 you(d make sure that the (ancestral fossils( to %hich you no% compared Trie =
%ere &raMBN1 &nd if Trie < happens to e &raMBN0 the (fossils( that you(d choose to unfreeFe and mix
them %ith %ould all e &raO1 The &raO and &raMBN !enes themselves0 as the -enski team already
kne% from previous %ork0 have no effect on fitness1 So they could use the colour markers to assay
the competitive ailities of each of the evolvin! tries0 usin! fossiliFed (ancestors( as the competitive
standard0 in every case1 &ll they had to do %as simply plate out samples from the mixed flasks and
see ho% many of the acteria !ro%in! on the a!ar %ere %hite and ho% many red1
&s # say0 in all t%elve tries the avera!e fitness increased as the thousands of !enerations
%ent y1 &ll t%elve lines !ot etter at survivin! in these !lucose:limited conditions1 The fitness
increase could e attriuted to several chan!es1 Populations !re% faster in successive flasks0 and the
avera!e ody siFe of the acteria !re%0 in all t%elve lines1 The top !raph opposite plots the avera!e
acterial ody siFe for one of the tries0 %hich %as typical1 The los represent real data points1 The
curve dra%n is a mathematical approximation1 #t !ives the est fit to the oserved data for this
particular kind of curve0 %hich is called a hyperola1L #t is al%ays possile that a more complicated
mathematical function than a hyperola %ould !ive an even closer fit to the data0 ut this hyperola
is pretty !ood0 so it hardly seems %orth otherin! to try1 Biolo!ists often fit mathematical curves to
oserved data0 ut0 unlike physicists0 iolo!ists are not accustomed to seein! such a close fit1
$sually our data are too messy1 #n iolo!y0 as opposed to physical sciences0 %e only expect to !et
smooth curves %hen %e have a very lar!e Euantity of data !athered under scrupulously controlled
conditions1 -enski(s research is a class act1
1ens&i e/#eri"ent: *aterial *o%y si-e in one tri*e
6ou can see that most of the increase in ody siFe occurred in the first 30555 or so
!enerations1 The next interestin! Euestion is this1 Given that all t%elve tries increased in ody siFe
over evolutionary time0 did they all increase in the same %ay0 y the same !enetic routeB *o0 they
didn(t0 and that(s the second interestin! result1 The !raph at the top of pa!e 234 is for one of the
t%elve tries1 *o% look at the eEuivalent hyperolic est fits for all t%elve 7!raph at the foot of
pa!e 23481 -ook ho% spread out they are1 They all seem to e approachin! a plateau0 ut the hi!hest
of the t%elve plateaus is almost t%ice as hi!h as the lo%est1 &nd the curves have different shapesA
the curve that reaches the hi!hest value y !eneration 250555 starts y !ro%in! more slo%ly than
some of the others0 and then overtakes them efore !eneration @05551 'on(t confuse these plateaus0
y the %ay0 %ith the daily plateaus of population siFe %ithin each flask1 We are no% lookin! at
curves in evolutionary time0 measured in flask !enerations0 not individual acterial time0 measured
in hours %ithin one flask1
1ens&i e/#eri"ent: *aterial *o%y si-e in t,el.e tri*es
What this evolutionary chan!e su!!ests is that ecomin! lar!er is0 for some reason0 a !ood
idea %hen you are stru!!lin! to survive in this alternatin! !lucose:richG!lucose:poor environment1 #
%on(t speculate on %hy increasin! ody siFe mi!ht e an advanta!e : there are many possiilities :
ut it looks as thou!h it must have een so0 ecause all t%elve tries did it1 But there are lots of
different %ays to ecome lar!er : different sets of mutations : and it looks as thou!h different %ays
have een discovered y different evolutionary linea!es in this experiment1 That(s pretty interestin!1
But perhaps even more interestin! is that sometimes a pair of tries seem to have independently
discovered the same %ay of !ettin! i!!er1 -enski and a different set of collea!ues investi!ated this
phenomenon y takin! t%o of the tries0 called &raO2 and &raMBN20 %hich seemed0 over 350555
!enerations0 to have follo%ed the same evolutionary traHectory0 and lookin! at their '*&1 The
astonishin! result they found %as that =9 !enes had chan!ed their levels of expression in oth
tries0 and all 78 had chan$ed in the same direction1 Were it not for natural selection0 such
independent parallelism0 in =9 !enes independently0 %ould completely e!!ar elief1 The odds
a!ainst its happenin! y chance are stupefyin!ly lar!e1 This is exactly the kind of thin! creationists
say cannot happen0 ecause they think it is too improale to have happened y chance1 6et it
actually happened1 &nd the explanation0 of course0 is that it did not happen y chance0 ut ecause
!radual0 step:y:step0 cumulative natural selection favoured the same : literally the same :
eneficial chan!es in oth lines independently1
1ens&i e/#eri"ent: inrease in $itness
The smooth curve in the !raph of increasin! cell siFe as the !enerations !o y !ives support
to the idea that the improvement is !radual1 But perhaps it is too !radualB Wouldn(t you expect to
see actual steps0 as the population (%aits( for the next improvin! mutation to turn upB *ot
necessarily1 #t depends on factors such as the numer of mutations involved0 the ma!nitude of each
mutation(s effect0 the variation in cell siFe that is caused y influences other than !enes0 and ho%
often the acteria %ere sampled1 &nd interestin!ly0 if %e look at the !raph of the increase in fitness0
as opposed to cell siFe0 %e do see %hat could at least e interpreted as a more overtly stepped
picture 7aove81 6ou rememer0 %hen # introduced the hyperola0 # said it mi!ht e possile to find
a more complicated mathematical function that %ould fit the data etter1 "athematicians call it a
(model(1 6ou could fit a hyperolic model to these points0 as in the previous !raph0 ut you !et an
even etter fit %ith a (step model(0 as used in this picture1 #t is not such a close fit as the cell siFe
!raph(s fit to a hyperola1 #n neither case can it e proved that the data exactly fit the model0 nor can
that ever e done1 But the data are at least compatile %ith the idea that the evolutionary chan!e
that %e oserve represents the step%ise accumulation of mutations1 L
We have so far seen a eautiful demonstration of evolution in actionA evolution efore our
very eyes0 documented y comparin! t%elve independent lines0 and also y comparin! each line
%ith (livin! fossils(0 %hich literally0 instead of only metaphorically0 come from the past1
*o% %e are ready to move on to an even more interestin! result1 So far0 #(ve implied that all
t%elve tries evolved their improved fitness in the same !eneral kind of %ay0 differin! only in detail
: some ein! a it faster0 some a it slo%er than others1 )o%ever0 the lon!:term experiment thre%
up one dramatic exception1 Shortly after !eneration 440555 somethin! utterly remarkale happened1
One out of the t%elve linea!es0 called &raMBN40 suddenly %ent erserk1 -ook at the !raph opposite1
The vertical axis0 laelled O'0 %hich stands for optical density or (cloudiness(0 is a measure of
population siFe in the flask1 The liEuid ecomes cloudy ecause of the sheer numers of acteriaJ
the thickness of the cloud can e measured as a numer0 and that numer is our index of population
density1 6ou can see that up to aout !eneration 4405550 the avera!e population density of Trie
&raMBN4 %as coastin! alon! at an O' of aout 515>0 %hich %as not very different from all the other
tries1 Then0 Hust after !eneration 4402550 the O' score of Trie &raMBN4 7and of that trie alone
amon! the t%elve8 %ent into vertical take:off1 #t shot up sixfold0 to an O' value of aout 513=1 The
populations of successive flasks of this trie soared1 &fter only a fe% days the typical plateau at
%hich flasks of this trie stailiFed had an O' numer aout six times !reater than it had een0 and
than the other tries %ere still sho%in!1 This hi!her plateau %as then reached in all suseEuent
!enerations0 in this trie ut no other1 #t %as as thou!h a lar!e dose of extra !lucose had een
inHected into every flask of Trie &raMBN40 ut !iven to no other trie1 But that didn(t happen1 The
same !lucose ration %as scrupulously administered to all the flasks eEually1
1ens&i e/#eri"ent: #o#(lation %ensity
What %as !oin! onB What %as it that suddenly happened to Trie &raMBN4B -enski and t%o
collea!ues investi!ated further0 and %orked it out1 #t is a fascinatin! story1 6ou rememer # said that
!lucose %as the limitin! resource0 and any mutant that (discovered( ho% to deal more efficiently
%ith !lucose %ould have an advanta!e1 That indeed is %hat happened in the evolution of all t%elve
tries1 But # also told you that !lucose %as not the only nutrient in the roth1 &nother one %as
citrate 7related to the sustance that makes lemons sour81 The roth contained plenty of citrate0 ut
E4 coli normally can(t use it0 at least not %here there is oxy!en in the %ater0 as there %as in -enski(s
flasks1 But if only a mutant could (discover( ho% to deal %ith citrate0 a onanFa %ould open up for
it1 This is exactly %hat happened %ith &raMBN41 This trie0 and this trie alone0 suddenly acEuired
the aility to eat citrate as %ell as !lucose0 rather than only !lucose1 The amount of availale food in
each successive flask in the linea!e therefore shot up1 &nd so did the plateau at %hich the
population in each successive flask daily stailiFed1
)avin! discovered %hat %as special aout the &raMBN4 trie0 -enski and his collea!ues %ent
on to ask an even more interestin! Euestion1 Was this sudden improvement in aility to dra%
nourishment all due to a sin!le dramatic mutation0 one so rare that only one of the t%elve linea!es
%as fortunate enou!h to under!o itB Was it0 in other %ords0 Hust another mutational step0 like the
ones that seemed to e demonstrated in the small steps of the fitness !raph on pa!e 23=B This
seemed to -enski unlikely0 for an interestin! reason1 Dno%in! the avera!e mutation rate of each
!ene in the !enome of these acteria0 he calculated that 450555 !enerations %as lon! enou!h for
every !ene to have mutated at least once in each of the t%elve lines1 So it seemed unlikely that it
%as the rarity of the mutation that sin!led &raMBN4 out1 #t should have een (discovered( y several
other tries1
There %as another theoretical possiility0 and an extremely tantaliFin! one1 This is %here
the story starts to !et Euite complicated so0 if it is late at ni!ht0 it mi!ht e an idea to resume readin!
tomorro% 1 1 1
What if the necessary iochemical %iFardry to feed on citrate reEuires not Hust one mutation
ut t%o 7or three8B We are not no% talkin! aout t%o mutations that uild on each other in a simple
additive %ay1 #f %e %ere0 it %ould e enou!h to !et the t%o mutations in any order1 Either one0 on
its o%n0 %ould take you half%ay 7say8 to the !oalJ and either one on its o%n %ould confer an aility
to !et some nourishment from citrate0 ut not as much as oth mutations to!ether %ould1 That
%ould e on a par %ith the mutations %e have already discussed for increasin! ody siFe1 But such
a circumstance %ould not e rare enou!h to account for the dramatic uniEueness of Trie &raMBN41
*o0 the rarity of citrate metaolism su!!ests that %e are lookin! for somethin! more like the
(irreducile complexity( of creationist propa!anda1 This mi!ht e a iochemical path%ay in %hich
the product of one chemical reaction feeds into a second chemical reaction0 and neither can make
any inroads at all witho"tthe other1 This %ould reEuire t%o mutations0 call them & and B0 to
catalyse the t%o reactions1 On this hypothesis0 you really %ould need both mutations before there is
any improement whatsoeer0 and that really %ould e improale enou!h to account for the
oserved result that only one out of the t%elve tries achieved the feat1
That(s all hypothetical1 Could the -enski !roup find out y experiment %hat %as actually
!oin! onB Well0 they could take !reat strides in that direction0 makin! rilliant use of the froFen
(fossils(0 %hich are such a continual oon in this research1 The hypothesis0 to repeat0 is that0 at some
time unkno%n0 Trie &raMBN4 chanced to under!o a mutation0 mutation &1 This had no detectale
effect ecause the other necessary mutation0 B0 %as still lackin!1 "utation B is eEually likely to
crop up in any one of the t%elve tries1 #ndeed0 it proaly did1 But B is no use : has asolutely no
eneficial effect at all : unless the trie happens to e primed y the previous occurrence of
mutation &1 &nd only trie &raMBN40 as it happened0 %as so primed1
-enski could even have phrased his hypothesis in the form of a testale prediction : and it is
interestin! to put it like this ecause it really is a prediction even thou!h0 in a sense0 it is aout the
past1 )ere(s ho% # %ould have put the prediction0 if # had een -enskiA
# shall tha% out fossils from Trie &ra:40 datin! from various points0 strate!ically chosen0
!oin! ack in time1 Each of these (-aFarus clones( %ill then e allo%ed to evolve further0 on a
similar re!imen to the main evolution experiment0 from %hich0 of course0 they %ill e completely
isolated1 &nd no%0 here(s my prediction1 Some of these -aFarus clones %ill (discover( ho% to deal
%ith citrate0 ut only if they %ere tha%ed out of the fossil record after a particular0 critical
!eneration in the ori!inal evolution experiment1 We don(t kno% : yet : %hen that ma!ic !eneration
%as ut %e shall identify it0 %ith hindsi!ht0 as the moment %hen0 accordin! to our hypothesis0
mutation & entered the trie1
6ou %ill e deli!hted to hear that this is exactly %hat -enski(s student Iachary Blount
found0 %hen he ran a !ruellin! set of experiments involvin! some forty trillion :
>50555055505550555 : E4 coli cells from across the !enerations1 The ma!ic moment turned out to e
approximately !eneration 3505551 Tha%ed:out clones of &raMBN4 that dated from after !eneration
350555 in the (fossil record( sho%ed increased proaility of s"bse3"ently evolvin! citrate
capaility1 *o clones that dated from efore !eneration 350555 did1 &ccordin! to the hypothesis0
after !eneration 350555 the clones %ere no% (primed( to take advanta!e of mutation B %henever it
came alon!1 &nd there %as no suseEuent chan!e in likelihood0 in either direction0 once the fossils(
(resurrection day( %as later than the ma!ic date of !eneration 350555A %hichever !eneration after
350555 Blount sampled0 the increased likelihood of those tha%ed fossils suseEuently acEuirin!
citrate capaility remained the same1 But tha%ed fossils from efore !eneration 350555 had no
increased likelihood of developin! citrate capaility at all1 Trie &raMBN40 efore !eneration 3505550
%as Hust like all the other tries1 &lthou!h its memers elon!ed to Trie &raMBN40 they did not
possess mutation &1 But after !eneration 3505550 Trie &raMBN4 %ere (primed(1 Only they %ere ale
to take advanta!e of (mutation B( %hen it turned up : as it proaly did in several of the other tries0
ut to no !ood effect1 There are moments of !reat Hoy in scientific research0 and this must surely
have een one of them1
-enski(s research sho%s0 in microcosm and in the la0 massively speeded up so that it
happened efore our very eyes0 many of the essential components of evolution y natural selectionA
random mutation follo%ed y non:random natural selectionJ adaptation to the same environment y
separate routes independentlyJ the %ay successive mutations uild on their predecessors to produce
evolutionary chan!eJ the %ay some !enes rely0 for their effects0 on the presence of other !enes1 6et
it all happened in a tiny fraction of the time evolution normally takes1
There is a comic seEuel to this triumphant tale of scientific endeavour1 Creationists hate it1
*ot only does it sho% evolution in actionJ not only does it sho% ne% information enterin! !enomes
%ithout the intervention of a desi!ner0 %hich is somethin! they have all een told to deny is
possile 7(told to( ecause most of them don(t understand %hat (information( means8J not only does it
demonstrate the po%er of natural selection to put to!ether cominations of !enes that0 y the naive
calculations so eloved of creationists0 should e tantamount to impossileJ it also undermines their
central do!ma of (irreducile complexity(1 So it is no %onder they are disconcerted y the -enski
research0 and ea!er to find fault %ith it1
&ndre% Schlafly0 creationist editor of (Conservapedia(0 the notoriously misleadin! imitation
of Wikipedia0 %rote to 'r -enski demandin! access to his ori!inal data0 presumaly implyin! that
there %as some dout as to their veracity1 -enski had asolutely no oli!ation even to reply to this
impertinent su!!estion ut0 in a very !entlemanly %ay0 he did so0 mildly su!!estin! that Schlafly
mi!ht make the effort to read his paper efore criticiFin! it1 -enski %ent on to make the tellin!
point that his est data are stored in the form of froFen acterial cultures0 %hich anyody could0 in
principle0 examine to verify his conclusions1 )e %ould e happy to send samples to any
acteriolo!ist Eualified to handle them0 pointin! out that in unEualified hands they mi!ht e Euite
dan!erous1 -enski listed these Eualifications in merciless detail0 and one can almost hear the relish
%ith %hich he did so0 kno%in! full %ell that Schlafly : a lawyer0 if you please0 not a scientist at all :
%ould hardly e ale to spell his %ay throu!h the %ords0 let alone Eualify as a acteriolo!ist
competent to carry out advanced and safe laoratory procedures0 follo%ed y statistical analysis of
the results1 The %hole matter %as trenchantly summed up y the celerated scientific lo!%it PI
"yers0 in a passa!e e!innin!0 (Once a!ain0 Richard -enski has replied to the !oons and fools at
Conservapedia0 and oy0 does he ever outclass them1(
-enski(s experiments0 especially %ith the in!enious (fossiliFation( techniEue0 sho% the po%er
of natural selection to %reak evolutionary chan!e on a timescale that %e can appreciate in a human
lifetime0 efore our very eyes1 But acteria provide other impressive0 if less clearly %orked:out0
examples1 "any acterial strains have evolved resistance to antiiotics in spectacularly short
periods1 &fter all0 the first antiiotic0 penicillin0 %as developed0 heroically0 y ,lorey and Chain as
recently as the Second World War1 *e% antiiotics have een comin! out at freEuent intervals since
then0 and acteria have evolved resistance to Hust aout every one of them1 *o%adays0 the most
ominous example is "RS& 7methycillin:resistant Staphylococc"s a"re"s80 %hich has succeeded in
makin! many hospitals positively dan!erous places to visit1 &nother menace is ( #4 diff4( 7
#lostridi"m difficile81 )ere a!ain0 %e have natural selection favourin! strains that are resistant to
antiioticsJ ut the effect is overlain y another one1 Prolon!ed use of antiiotics tends to kill (!ood(
acteria in the !ut0 alon! %ith the ad ones1 #4 diff40 ein! resistant to most antiiotics0 is !reatly
helped y the absence of other species of acteria %ith %hich it %ould normally compete1 #t is the
principle of (my enemy(s enemy is my friend(1
# %as mildly irritated to read a pamphlet in my doctor(s %aitin! room %arnin! of the dan!er
of failin! to finish a course of antiiotic pills1 *othin! %ron! %ith that %arnin!J ut it %as the
reason !iven that %orried me1 The pamphlet explained that acteria are (clever(J they (learn( to cope
%ith antiiotics1 Presumaly the authors thou!ht the phenomenon of antiiotic resistance %ould e
easier to !rasp if they called it learnin! rather than natural selection1 But to talk of acteria ein!
clever0 and of learnin!0 is do%nri!ht confusin!0 and aove all it doesn(t help the patient to make
sense of the instruction to carry on takin! the pills until they are finished1 &ny fool can see that it is
not plausile to descrie a acterium as clever1 Even if there %ere clever acteria0 %hy %ould
stoppin! prematurely make any difference to the learnin! pro%ess of a clever acteriumB But as
soon as you start thinkin! in terms of natural selection0 it makes perfect sense1
-ike any poison0 antiiotics are likely to e dosa!e dependent1 & sufficiently hi!h dose %ill
kill all the acteria1 & sufficiently lo% dose %ill kill none1 &n intermediate dose %ill kill some0 ut
not all1 #f there is !enetic variation amon! acteria0 such that some are more susceptile to the
antiiotic than others0 an intermediate dose %ill e tailor:made to select in favour of !enes for
resistance1 When the doctor tells you to finish takin! the pills0 it is to increase the chances of killin!
all the acteria and avoid leavin! ehind resistant0 or semi:resistant0 mutants1 With hindsi!ht %e
mi!ht say that if only %e had all een etter educated in 'ar%inian thinkin!0 %e %ould have %oken
up sooner to the dan!ers of resistant strains ein! selected1 Pamphlets like the one in my doctor(s
%aitin! room don(t help %ith that education : and %hat a sadly missed opportunity to teach
somethin! of the %ondrous po%er of natural selection1
"y collea!ue 'r .ohn Endler0 recently moved from *orth &merica to the $niversity of
Exeter0 told me the follo%in! marvellous : %ell0 also depressin! : story1 )e %as travellin! on a
domestic fli!ht in the $nited States0 and the passen!er in the next seat made conversation y askin!
him %hat he did1 Endler replied that he %as a professor of iolo!y0 doin! research on %ild !uppy
populations in Trinidad1 The man ecame increasin!ly interested in the research and asked many
Euestions1 #ntri!ued y the ele!ance of the theory that seemed to underlie the experiments0 he asked
Endler %hat that theory %as0 and %ho ori!inated it1 Only then did 'r Endler drop %hat he correctly
!uessed %ould e his omshellA (#t(s called 'ar%in(s theory of evolution y natural selectionK( The
man(s %hole demeanour instantly chan!ed1 )is face %ent redJ aruptly0 he turned a%ay0 refused to
speak further and terminated %hat had hitherto een an amiale conversation1 "ore than amiale0
indeedA 'r Endler %rites to me that the man had (asked some excellent Euestions efore this0
indicatin! that he %as enthusiastically and intellectually follo%in! the ar!ument1 This is really
The experiments that .ohn Endler recounted to his closed:minded fello% passen!er are
ele!ant and simple0 and they serve eautifully to illustrate the speed %ith %hich natural selection
can !o to %ork1 #t is fittin! that # should use Endler(s o%n research here0 ecause he is also the
author of .at"ral Selection in the Wild0 the leadin! ook in %hich examples of such studies have
een collected0 and their methods laid out1
Guppies are popular fresh%ater aEuarium fish1 &s %ith the pheasants %e met in Chapter 40
the males are more ri!htly coloured than the females0 and aEuarists have red them to ecome
even ri!hter1 Endler studied %ild !uppies 7Poecilia retic"lata8 livin! in mountain streams in
Trinidad0 Toa!o and +eneFuela1 )e noticed that local populations %ere strikin!ly different from
each other1 #n some populations the adult males %ere raino%:coloured0 almost as ri!ht as those
red in aEuarium tanks1 )e surmised that their ancestors had een selected for their ri!ht colours
y female !uppies0 in the same manner as cock pheasants are selected y hens1 #n other areas the
males %ere much draer0 althou!h they %ere still ri!hter than the females1 -ike the females0
thou!h less so0 they %ere %ell camoufla!ed a!ainst the !ravelly ottoms of the streams in %hich
they live1 Endler sho%ed0 y ele!ant Euantitative comparisons et%een many locations in
+eneFuela and Trinidad0 that the streams %here the males %ere less ri!ht %ere also the streams
%here predation %as heavy1 #n streams %ith only %eak predation0 males %ere more ri!htly
coloured0 %ith lar!er0 !audier spots0 and more of themA here the males %ere free to evolve ri!ht
colours to appeal to females1 The pressure from females on males to evolve ri!ht colours %as there
all the time0 in all the various separate populations0 %hether the local predators %ere pushin! in the
other direction stron!ly or %eakly1 &s ever0 evolution finds a compromise et%een selection
pressures1 What %as interestin! aout the !uppies is that Endler could actually see ho% the
compromise varied in different streams1 But he did much etter than that1 )e %ent on to do
Suppose you %anted to set up the ideal experiment to demonstrate the evolution of
camoufla!eA %hat %ould you doB Camoufla!ed animals resemle the ack!round on %hich they are
seen1 Could you set up an experiment in %hich animals actually evolve0 efore your very eyes0 to
resemle a ack!round that you have experimentally provided for themB Preferaly t%o
ack!rounds0 %ith a different population on eachB The aim is to do somethin! like the selection of
t%o lines of maiFe plants for hi!h and lo% oil content that %e sa% in Chapter 41 But in these
experiments the selection %ill e done not y humans ut y predators and y female !uppies1 The
only thin! that %ill separate the t%o experimental lines is the different ack!rounds that %e shall
Take some animals of a camoufla!ed species0 perhaps a species of insect0 and assi!n them
randomly to different ca!es 7or enclosures0 or ponds0 %hatever is suitale8 %hich have differently
coloured0 or differently patterned0 ack!rounds1 ,or example0 you mi!ht !ive half the enclosures a
!reen foresty ack!round and the other half a reddish:ro%n0 deserty ack!round1 )avin! put your
animals in their !reen or ro%n enclosures0 you(d then leave them to live and reed for as many
!enerations as you have time for0 after %hich you(d come ack to see %hether they had evolved to
resemle their ack!rounds0 !reen or ro%n respectively1 Of course0 you only expect this result if
you put predators in the enclosure too1 So0 let(s put0 say0 a chameleon in1 #n all the enclosuresB *o0
of course not1 This is an experiment0 rememerJ so you(d put a predator in half the !reen enclosures
and half the red enclosures1 The experiment %ould e to test the prediction that0 in enclosures %ith a
predator0 the insects %ould evolve to ecome either !reen or ro%n : to ecome more similar to
their ack!round1 But in the enclosures %ithout a predator0 they mi!ht if anythin! evolve to ecome
more different from their ack!round0 to e conspicuous to females1
# have lon! nursed an amition to do exactly this experiment %ith fruit flies 7ecause their
reproductive turnover time is so short8 ut0 alas0 # never !ot around to it1 So # am especially
deli!hted to say that this is exactly %hat .ohn Endler did0 not %ith insects ut %ith !uppies1
Oviously he didn(t use chameleons for predators0 ut instead chose a fish called the pike cichlid
7pronounced (sick lid(80 #renicichla alta0 %hich is a dan!erous predator of these !uppies in the %ild1
*or did he use !reen versus ro%n ack!rounds : he opted for somethin! more interestin! than that1
)e noticed that !uppies derive much of their camoufla!e from their spots0 often Euite lar!e ones0
%hose patternin! resemles the patternin! of the !ravelly ottoms of their native streams1 Some
streams have coarser0 more pely !ravel0 others finer0 more sandy !ravel1 Those %ere the t%o
ack!rounds he used0 and you(ll a!ree that the camoufla!e he %as seekin! %as sutler and more
interestin! than my !reen versus ro%n1
Endler !ot a lar!e !reenhouse0 to simulate the tropical %orld of the !uppies0 and set up ten
ponds inside it1 )e put !ravel on the ottom of all ten ponds0 ut five of them had coarse0 pely
!ravel and the other five had finer0 sandy !ravel1 6ou can see %here this is !oin!1 The prediction is
that0 %hen exposed to stron! predation0 the !uppies on the t%o ack!rounds %ill diver!e from each
other over evolutionary time0 each in the direction of matchin! its o%n ack!round1 Where
predation is %eak or non:existent0 the prediction is that the males should tend in the direction of
ecomin! more conspicuous0 to appeal to females1
#nstead of puttin! predators in half the ponds and no predators in the other half0 a!ain Endler
did somethin! more sutle1 )e had three levels of predation1 T%o ponds 7one fine and one coarse
!ravel8 had no predators at all1 ,our ponds 7t%o fine and t%o coarse !ravel8 had the dan!erous pike
cichlid1 #n the remainin! four ponds0 Endler introduced another species of fish0 Ri"l"s hartii0
%hich0 despite its En!lish name0 (killifish( 7actually that(s Euite irrelevant since it is named after a
"r Dille80 is relatively harmless to !uppies1 #t is a (%eak predator(0 %hereas the pike cichlid is a
stron! predator1 The (%eak predator( situation is a etter control condition than no predators at all1
This is ecause0 as Endler explains0 he %as tryin! to simulate t%o natural conditions0 and he kno%s
of no natural streams that are totally free of predatorsA thus the comparison et%een stron! and
%eak predation is a more natural comparison1
So0 here(s the set:upA !uppies %ere assi!ned randomly to ten ponds0 five %ith coarse !ravel
and five %ith fine !ravel1 &ll ten colonies of !uppies %ere allo%ed to reed freely for six months
%ith no predators1 &t this point the experiment proper e!an1 Endler put one (dan!erous predator(
into each of t%o coarse !ravel ponds and t%o fine !ravel ponds1 )e put six (%eak predators( 7six
rather than one0 to !ive a closer approximation to the relative densities of the t%o kinds of fish in
the %ild8 into each of t%o coarse !ravel ponds and t%o fine !ravel ponds1 &nd the remainin! t%o
ponds Hust carried on as efore0 %ith no predators at all1
&fter the experiment had een runnin! for five months0 Endler took a census of all the
ponds0 and counted and measured the spots on all the !uppies in all the ponds1 *ine months later0
that is0 after fourteen months in all0 he took another census0 countin! and measurin! in the same
%ay1 &nd %hat of the resultsB They %ere spectacular0 even after so short a time1 Endler used
various measures of the fishes( colour patterns0 one of %hich %as (spots per fish(1 When the !uppies
%ere first put into their ponds0 efore the predators %ere introduced0 there %as a very lar!e ran!e of
spot numers0 ecause the fish had een !athered from a %ide variety of streams0 of %idely varyin!
predator content1 'urin! the six months efore any predators %ere introduced0 the mean numer of
spots per fish shot up1 Presumaly this %as in response to selection y females1 Then0 at the point
%hen the predators %ere introduced0 there %as a dramatic chan!e1 #n the four ponds that had the
dan!erous predator0 the mean numer of spots plummeted1 The difference %as fully apparent at the
five:month census0 and the numer of spots had declined even further y the fourteen:month
census1 But in the t%o ponds %ith no predators0 and the four ponds %ith %eak predation0 the
numer of spots continued to increase1 #t reached a plateau as early as the five:month census0 and
stayed hi!h for the fourteen:month census1 With respect to spot numer0 %eak predation seems to
e pretty much the same as no predation0 over:ruled y sexual selection y females %ho prefer lots
of spots1
So much for spot numer1 Spot siFe tells an eEually interestin! story1 #n the presence of
predators0 %hether %eak or stron!0 coarse !ravel promoted relatively lar!er spots0 %hile fine !ravel
favoured relatively smaller spots1 This is easily interpreted as spot siFe mimickin! stone siFe1
,ascinatin!ly0 ho%ever0 in the ponds %here there %ere no predators at all0 Endler found exactly the
reverse1 ,ine !ravel favoured lar!e spots on male !uppies0 and coarse !ravel favoured small spots1
They are more conspicuous if they do not mimic the stones on their respective ack!rounds0 and
that is !ood for attractin! females1 *eatK
6es0 neat1 But that %as in the la1 Could Endler !et similar results in the %ildB 6es1 )e %ent
to a natural stream that contained the dan!erous pike cichlids0 in %hich the male !uppies %ere all
relatively inconspicuous1 )e cau!ht !uppies of oth sexes and transplanted them to a triutary of
the same stream that contained no !uppies and no dan!erous predators0 althou!h the %eak predator
killifish %ere present1 )e left them there to !et on %ith livin! and reedin!0 and %ent a%ay1
T%enty:three months later0 he returned and re:examined the !uppies to see %hat had happened1
&maFin!ly0 after less than t%o years0 the males had shifted noticealy in the direction of ein! more
ri!htly coloured : pulled y females0 no dout0 and freed to !o there y the asence of dan!erous
One of the nice thin!s aout science is that it is a pulic activity1 Scientists pulish their
methods as %ell as their conclusions0 %hich means that anyody else0 any%here in the %orld0 can
repeat their %ork1 #f they don(t !et the same results0 %e %ant to kno% the reason %hy1 $sually they
don(t Hust repeat previous %ork ut extend itA carry it further1 .ohn Endler(s rilliant research on
!uppies %as Hust e!!in! to e continued and extended1 &mon! those %ho have taken it up is 'avid
ReFnick of the $niversity of California at Riverside1
*ine years after Endler sampled his experimental stream %ith such spectacular results0
ReFnick and his collea!ues revisited the place and sampled the descendants of Endler(s
experimental population yet a!ain1 The males %ere no% very ri!htly coloured1 The female:driven
trend that Endler oserved had continued0 %ith a ven!eance1 &nd that %asn(t all1 6ou rememer the
silver foxes of Chapter 40 and ho% artificial selection for one characteristic 7tameness8 pulled alon!
in its %ake a %hole cluster of othersA chan!es in reedin! season0 in ears0 tail0 coat colour and other
thin!sB Well0 a similar thin! happened %ith the !uppies0 under natural selection1
ReFnick and Endler had already noticed that %hen you compare !uppies in predator:infested
streams %ith !uppies in streams %ith only %eak predation0 colour differences are only the tip of the
iceer!1 There is a %hole cluster of other differences1 Guppies from lo%:predation streams reach
sexual maturity later than those from hi!h:predation streams0 and they are lar!er %hen they reach
adulthoodJ they produce litters of youn! less freEuentlyJ and their litters are smaller0 %ith lar!er
offsprin!1 When ReFnick examined the descendants of Endler(s !uppies0 his findin!s %ere almost
too !ood to e true1 The ones that had een freed to follo% female:driven sexual selection rather
than predator:driven selection for individual survival had not only ecome more ri!htly colouredA
in all the other respects # have Hust listed0 these fish had evolved the full cluster of other chan!es0 to
match those normally found in %ild populations free from predators1 The !uppies matured at a later
a!e than in predator:infested streams0 they %ere lar!er0 and they produced fe%er and lar!er
offsprin!1 The alance had shifted to%ards the norm for predator:free pools0 %here sexual
attractiveness takes priority1 &nd it all happened sta!!erin!ly fast0 y evolutionary standards1 -ater
in the ook %e shall see that the evolutionary chan!e %itnessed y Endler and ReFnick0 driven
purely y natural selection 7strictly includin! sexual selection80 raced ahead at a speed comparale
to that achieved y artificial selection of domestic animals1 #t is a spectacular example of evolution
efore our very eyes1
One of the surprisin! thin!s %e have learned aout evolution is that it can e oth very fast :
as %e have seen in this chapter : and0 under other circumstances0 as %e kno% from the fossil record0
very slo%1 Slo%est of all are those livin! creatures that %e call (livin! fossils(1 They are not literally
rou!ht ack from the dead like -enski(s froFen acteria1 But they are creatures that have chan!ed
so little since their remote ancestors that it is almost as thou!h they %ere fossils1
"y favourite livin! fossil is the rachiopod (in$"la4 6ou don(t need to kno% %hat a
rachiopod is1 They %ould surely have een staples on the menu0 had seafood restaurants flourished
efore the !reat Permian extinction a Euarter of a illion years a!o : the most catastrophic extinction
of all time1 & superficial !lance mi!ht confuse them %ith ivalve molluscs : mussels and their kind
: ut they are really very different1 Their t%o shells are top and ottom0 %here mussels( shells are
left and ri!ht1 #n evolutionary history ivalves and rachiopods %ere0 as Stephen .ay Gould
memoraly put it0 ships that pass in the ni!ht1 & fe% rachiopods survived (the Great 'yin!(
7Gould(s phrase a!ain80 and modern (in$"la 7aove8 is so similar to (in$"lella0 the fossil elo%0
that the fossil %as ori!inally !iven the same !eneric name0 (in$"la1 This particular specimen of
(in$"lella !oes ack to the Ordovician era0 >=5 million years a!o1 But there are fossils0 also
ori!inally named (in$"la and no% kno%n as (in$"lella0 !oin! ack more than half a illion years
to the Camrian era1 # should admit0 ho%ever0 that a fossiliFed shell is not a lot to !o on0 and some
Foolo!ists dispute (in$"la(s claim to e an almost %holly unchan!ed (livin! fossil(1
Lingulella ) al"ost i%ential to its "o%ern
"any of the prolems that %e meet in evolutionary ar!umentation arise only ecause
animals are inconsiderate enou!h to evolve at different rates0 and mi!ht even e inconsiderate
enou!h not to evolve at all1 #f there %ere a la% of nature dictatin! that Euantity of evolutionary
chan!e must al%ays e oli!in!ly proportional to elapsed time0 de!ree of resemlance %ould
faithfully reflect closeness of cousinship1 #n the real %orld0 ho%ever0 %e have to contend %ith
evolutionary sprinters like irds0 %ho leave their reptile ori!ins standin! in the "esoFoic dust :
helped0 in our perception of their uniEueness0 y the happenstance that their nei!hours in the
evolutionary tree %ere all killed y a celestial catastrophe1 &t the other extreme0 %e have to contend
%ith (livin! fossils( like (in$"la %hich0 in extreme cases0 have chan!ed so little that they mi!ht
almost interreed %ith their remote ancestors0 if only a matchmakin! time:machine could procure
them a date1
(in$"la is not the only famous example of a livin! fossil1 Others include (im"l"s0 the
horseshoe (cra(0 and coelacanths0 %hich %e shall meet in the next chapter1
L Think of it like this1 #ma!ine all possile strai!ht lines1 ,or each line0 calculate ho%
closely it fits the dots0 y measurin! the distance of each dot from the line0 and addin! up all the
distances 7after sEuarin! them0 for a !ood mathematical reason %hich %ould take us too far afield81
Of all possile strai!ht lines0 the one that minimi2es the sum of the sEuared dot:to:line distances0
summed over all the dots0 is the fitted re!ression line1 #t sho%s us the trend0 %ithout our eyes ein!
distracted y all the mess of the individual points1 There are separate calculations that statisticians
do to calculate ho% reliable the line is as an indicator of a trend1 These are called tests of statistical
si!nificance1 They make use of the readth of scatter aout the line1
L $p to t%ice as lon! if the Pod Dopiste liFards have een evolvin! at the same rate since
the shared ancestor of thirty:seven years a!o1
L The same !orilla:like features in the skull and teeth of our roust cousin Paranthrop"s
boisei 7(nutcracker man(0 also nicknamed (IinH( and ('ear Boy(8 indicate that it %as almost certainly
L That(s no idle metaphor0 for fairy rin!s of mushrooms achieve their circular form for
exactly the same reason1
L 6ou rememer the strai!ht line that %as the est fit to the data on the decline of elephant
tusk siFe from 293= to 29=;B # explained the method as eEuivalent to tryin! all possile strai!ht
lines and findin! the one that minimiFed the sum of sEuares of distances of dots on the !raph from
the line1 But you can do the same thin! %hile not limitin! yourself to strai!ht lines1 6ou can look at
all possile curves of a certain type defined y mathematicians1 The hyperola is one such curve1 #n
this case0 %e look at all possile hyperolas in turn0 measure the distance to the line of every point
on the !raph0 then add up the sum of the sEuared distances over all points1 'o the same for all
hyperolas0 and then choose the hyperola that minimiFes that sum1 -enski did a sort of short:cut
eEuivalent to that exhaustive operation to arrive at the est:fit hyperola0 %hich is the one you see
dra%n in1
L & step%ise pattern of evolution is to e expected in creatures such as acteria0 %hich
7most of the time8 don(t reproduce sexually1 #n animals like us0 %ho reproduce only sexually0
evolutionary chan!e doesn(t usually (han! aout( %hile it (%aits( for a key mutation to turn up 7this is
a common mistake made y opponents of evolution %ith some pretensions to sophistication81
#nstead0 sexually reproducin! populations usually have a ready supply of !enetic variation from
%hich to select1 &lthou!h ori!inally !enerated y past mutation0 a lar!e numer of !enetic variants
are often present in a !ene pool at any one time0 introduced y mutation a %hile ack and no%
shuffled aout y sexual recomination1 *atural selection often acts to shift the alance of existin!
variation0 rather than %aitin! for key mutations to turn up1 #n acteria %ithout sexual reproduction0
the very idea of a !ene pool doesn(t properly apply1 That is %hy %e can realistically hope to see
discrete steps0 %here %e mi!ht not in a population of irds0 mammals or fish1
"#SS#*G -#*DB W)&T 'O 6O$ "E&*0 ("#SS#*G(B
C RE&T#O*#STS are deeply enamoured of the fossil record0 ecause they have een tau!ht
7y each other8 to repeat0 over and over0 the mantra that it is full of (!aps(A (Sho% me your
QintermediatesQK( They fondly 7very fondly8 ima!ine that these (!aps( are an emarrassment to
evolutionists1 &ctually0 %e are lucky to have any fossils at all0 let alone the massive numers that
%e no% do have to document evolutionary history : lar!e numers of %hich0 y any standards0
constitute eautiful (intermediates(1 # shall emphasiFe in Chapters 9 and 25 that %e don(t need fossils
in order to demonstrate that evolution is a fact1 The evidence for evolution %ould e entirely secure0
even if not a sin!le corpse had ever fossiliFed1 #t is a onus that %e do actually have rich seams of
fossils to mine0 and more are discovered every day1 The fossil evidence for evolution in many maHor
animal !roups is %onderfully stron!1 *evertheless there are0 of course0 !aps0 and creationists love
them osessively1
-et(s a!ain make use of our analo!y of the detective comin! to the scene of a crime to %hich
there %ere no eye %itnesses1 The aronet has een shot1 ,in!erprints0 footprints0 '*& from a s%eat
stain on the pistol0 and a stron! motive all point to%ards the utler1 #t(s pretty much an open and
shut case0 and the Hury and everyody in the court is convinced that the utler did it1 But a last:
minute piece of evidence is discovered0 in the nick of time efore the Hury retires to consider %hat
had seemed to e their inevitale verdict of !uiltyA someody rememers that the aronet had
installed spy cameras a!ainst ur!lars1 With ated reath0 the court %atches the films1 One of them
sho%s the utler in the act of openin! the dra%er in his pantry0 takin! out a pistol0 loadin! it0 and
creepin! stealthily out of the room %ith a malevolent !leam in his eye1 6ou mi!ht think that this
solidifies the case a!ainst the utler even further1 "ark the seEuel0 ho%ever1 The utler(s defence
la%yer astutely points out that there %as no spy camera in the lirary %here the murder took place0
and no spy camera in the corridor leadin! from the utler(s pantry1 )e %a!s his fin!er0 in that
compellin! %ay that la%yers have made their o%n1 (There(s a $ap in the video recordK We don(t
kno% %hat happened after the utler left the pantry1 There is clearly insufficient evidence to convict
my client1(
#n vain the prosecution la%yer points out that there %as a second camera in the illiard
room0 and this sho%s0 throu!h the open door0 the utler0 !un at the ready0 creepin! on tiptoe alon!
the passa!e to%ards the lirary1 Surely this plu!s the !ap in the video recordB Surely the case
a!ainst the utler is no% unassailaleB But no1 Triumphantly the defence la%yer plays his ace1 (We
don(t kno% %hat happened efore or after the utler passed the open door of the illiard room1
There are no% two !aps in the video record1 -adies and !entlemen of the Hury0 my case rests1 There
is no% even less evidence a!ainst my client than there %as efore1(
The fossil record0 like the spy camera in the murder story0 is a bon"s0 somethin! that %e had
no ri!ht to expect as a matter of entitlement1 There is already more than enou!h evidence to convict
the utler %ithout the spy camera0 and the Hury %ere aout to deliver a !uilty verdict efore the spy
camera %as discovered1 Similarly0 there is more than enou!h evidence for the fact of evolution in
the comparative study of modern species 7Chapter 258 and their !eo!raphical distriution 7Chapter
981 We don(t need fossils : the case for evolution is %aterti!ht %ithout themJ so it is paradoxical to
use $aps in the fossil record as thou!h they %ere evidence a!ainst evolution1 We are0 as # say0 lucky
to have fossils at all1
What wo"ld e evidence a!ainst evolution0 and very stron! evidence at that0 %ould e the
discovery of even a sin!le fossil in the %ron! !eolo!ical stratum1 # have already made this point in
Chapter >1 .1 B1 S1 )aldane famously retorted0 %hen asked to name an oservation that %ould
disprove the theory of evolution0 (,ossil raits in the PrecamrianK( *o such raits0 no
authentically anachronistic fossils of any kind0 have ever een found1 &ll the fossils that %e have0
and there are very very many indeed0 occur0 %ithout a sin!le authenticated exception0 in the ri!ht
temporal seEuence1 6es0 there are !aps0 %here there are no fossils at all0 and that is only to e
expected1 But not a sin!le solitary fossil has ever een found before it could have evolved1 That is a
very tellin! fact 7and there is no reason %hy %e should expect it on the creationist theory81 &s #
riefly mentioned in Chapter >0 a !ood theory0 a scientific theory0 is one that is vulnerale to
disproof0 yet is not disproved1 Evolution could so easily e disproved if Hust a sin!le fossil turned up
in the %ron! date order1 Evolution has passed this test %ith flyin! colours1 Sceptics of evolution
%ho %ish to prove their case should e dili!ently scralin! around in the rocks0 desperately tryin!
to find anachronistic fossils1 "aye they(ll find one1 Want a etB
The i!!est !ap0 and the one the creationists like est of all0 is the one that preceded the so:
called Camrian Explosion1 & little more than half a illion years a!o0 in the Camrian era0 most of
the !reat animal phyla : the main divisions %ithin the animal %orld : (suddenly( appear in the fossil
record1 Suddenly0 that is0 in the sense that no fossils of these animal !roups are kno%n in rocks
older than the Camrian0 not suddenly in the sense of instantaneouslyA the period %e are talkin!
aout covers aout 35 million years1 T%enty million years feels short %hen it is half a illion years
a!o1 But of course it represents exactly the same amount of time for evolution as 35 million years
todayK &ny%ay0 it is still Euite sudden0 and0 as # %rote in a previous ook0 the Camrian sho%s us a
sustantial numer of maHor animal phyla
already in an advanced state of evolution0 the very first time they appear1 #t is as thou!h they
%ere Hust planted there0 %ithout any evolutionary history1 *eedless to say0 this appearance of
sudden plantin! has deli!hted creationists1
That last sentence sho%s that # %as savvy enou!h to realiFe that creationists %ould like the
Camrian Explosion1 # %as not 7ack in 29;<8 savvy enou!h to realiFe that they(d !leefully Euote
my lines ack at me in their o%n favour0 over and over a!ain0 carefully omittin! my careful %ords
of explanation1 On a %him0 # Hust searched the World Wide We for (#t is as thou!h they %ere Hust
planted there0 %ithout any evolutionary history( and otained no fe%er than 203=5 hits1 &s a crude
control test of the hypothesis that the maHority of these hits represent creationist Euote:minin!s0 #
tried searchin!0 as a comparison0 for the clause that immediately follo%s the aove Euotation in The
Blind WatchmakerA (Evolutionists of all stripes elieve0 ho%ever0 that this really does represent a
very lar!e !ap in the fossil record(1 # otained a !rand total of <4 hits0 compared to the 203=5 hits for
the previous sentence1 The ratio of 23=5 to <4 is 291;1 We mi!ht call this ratio the Puote "inin!
# have dealt %ith the Camrian Explosion at len!th0 especially in 9nweain$ the Rainbow1
)ere #(ll add Hust one ne% point0 illustrated y the flat%orms0 Platyhelminthes1 This !reat phylum of
%orms includes the parasitic flukes and tape%orms0 %hich are of !reat medical importance1 "y
favourites0 ho%ever0 are the free:livin! turellarian %orms0 of %hich there are more than four
thousand speciesA that(s aout as numerous as all the mammal species put to!ether1 Some of these
turellarians are creatures of !reat eauty0 as the t%o pictured opposite sho%1 They are common0
oth in %ater and on land0 and presumaly have een common for a very lon! time1 6ou(d expect0
therefore0 to see a rich fossil history1 $nfortunately0 there is almost nothin!1 &part from a handful
of ami!uous trace fossils0 not a sin!le fossil flat%orm has ever een found1 The Platyhelminthes0
to a %orm0 are (already in an advanced state of evolution0 the very first time they appear1 #t is as
thou!h they %ere Hust planted there0 %ithout any evolutionary history1( But in this case0 (the very
first time they appear( is not the Camrian ut today1 'o you see %hat this means0 or at least ou!ht
to mean for creationistsB Creationists elieve that flat%orms %ere created in the same %eek as all
other creatures1 They have therefore had exactly the same time in %hich to fossiliFe as all other
animals1 'urin! all the centuries %hen all those ony or shelly animals %ere depositin! their fossils
y the thousands0 the flat%orms must have een livin! happily alon!side them0 ut %ithout leavin!
any si!nificant trace of their presence in the rocks1 What0 then0 is so special aout !aps in the record
of those animals that do fossiliFe0 !iven that the past history of the flat%orms amounts to one bi$
$apA even thou!h the flat%orms0 y the creationists( o%n account0 have een livin! for the same
len!th of timeB #f the !ap efore the Camrian Explosion is used as evidence that most animals
suddenly spran! into existence in the Camrian0 exactly the same (lo!ic( should e used to prove
that the flat%orms spran! into existence yesterday1 6et this contradicts the creationist(s elief that
flat%orms %ere created durin! the same creative %eek as everythin! else1 6ou cannot have it oth
%ays1 This ar!ument0 at a stroke0 completely destroys the creationist case that the Precamrian !ap
in the fossil record %eakens the evidence for evolution1
T(r*ellarians ) no $ossil reor%, *(t they "(st ha.e *een there all alon'
Why0 on the evolutionary vie%0 are there so fe% fossils efore the Camrian eraB Well0
presumaly0 %hatever factors applied to the flat%orms throu!hout !eolo!ical time to this day0 those
same factors applied to the rest of the animal kin!dom efore the Camrian1 Proaly0 most animals
efore the Camrian %ere soft:odied like modern flat%orms0 proaly also rather small like
modern turellarians : Hust not !ood fossil material1 Then somethin! happened half a illion years
a!o to allo% animals to fossiliFe freely : the arisin! of hard0 mineraliFed skeletons0 for example1
&n earlier name for (!ap in the fossil record( %as (missin! link(1 The phrase enHoyed a vo!ue
in late +ictorian En!land0 and persisted into the t%entieth century1 #nspired y a misunderstandin!
of 'ar%in(s theory0 it %as used as an insult in rou!hly the same %ay as (neanderthal( is colloEuially
7and unHustly8 used today1 &mon! the !xford En$lish Dictionary(s list of representative Euotations
is a 2945 one in %hich '1 )1 -a%rence tells of a %oman %ho %rote to say his name (stank( and %ent
on0 (6ou0 %ho are a mixture of the missin!:link and the chimpanFee1(
The ori!inal meanin!0 a confused one as # shall sho%0 implied that the 'ar%inian theory
lacked a vital link et%een humans and other primates1 &nother of the dictionary(s illustrative
Euotations0 a +ictorian one0 uses it like thisA (#(ve heard talk o( some missin! link0 at%een men and
pu!!ies( 7(pu!!ie( %as a Scottish dialect %ord for monkey81 )istory:deniers0 to this day0 are very
fond of sayin!0 in %hat they ima!ine is a tauntin! tone of voiceA (But you still haven(t found the
missin! link0( and they often thro% in a Hie aout Piltdo%n "an0 for !ood measure1 *oody kno%s
%ho perpetrated the Piltdo%n hoax0 ut he has a lot to ans%er for1L The fact that one of the first
candidates for a man:ape fossil to e discovered turned out to e a hoax provided an excuse for
history:deniers to i!nore the very numerous fossils that are not hoaxesJ and they still haven(t
stopped cro%in! aout it1 #f only they %ould look at the facts0 they(d soon discover that %e no%
have a rich supply of intermediate fossils linkin! modern humans to the common ancestor that %e
share %ith chimpanFees1 On the human side of the divide0 that is1 #nterestin!ly0 there are as yet no
fossils linkin! that ancestor 7%hich %as neither chimpanFee nor human8 to modern chimpanFees1
Perhaps this is ecause chimpanFees live in forests0 %hich don(t provide !ood fossiliFin! conditions1
#f anythin! it is chimpanFees0 not humans0 %ho today have a ri!ht to complain of missin! linksK
That0 then0 is one meanin! of (missin! link(1 #t is the alle!ed !ap et%een humans and the
rest of the animal kin!dom1 The missin! link in that sense is0 to put it mildly0 no lon!er missin!1 #
shall return to this in the next chapter0 %hich is specifically aout human fossils1
&nother meanin! concerns the alle!ed paucity of so:called (transitional forms( et%een
maHor !roupsA et%een reptiles and irds0 for example0 or et%een fish and amphiians1 (Produce
your intermediatesK( Evolutionists often respond to this challen!e from history:deniers y thro%in!
them the ones of Archaeopteryx0 the famous (intermediate( et%een (reptiles( and irds1 This is a
mistake0 as # shall sho%1 Archaeopteryx is not the ans%er to a challen!e0 ecause there is no
challen!e %orth ans%erin!1 To put up a sin!le famous fossil like Archaeopteryx panders to a
fallacy1 #n fact0 for a lar!e numer of fossils0 a !ood case can e made that every one of them is an
intermediate et%een somethin! and somethin! else1 The alle!ed challen!e that seems to e
ans%ered y Archaeopteryx is ased on an outdated conception0 the one that used to e kno%n as
the Great Chain of Bein!J and that is the title under %hich # shall deal %ith it later in this chapter1
The silliest of all these (missin! link( challen!es are the follo%in! t%o 7or variants of them0
of %hich there are many81 ,irst0 (#f people came from monkeys via fro!s and fish0 then %hy does the
fossil record not contain a QfronkeyQB( # have seen an #slamic creationist ask truculently %hy there
are no crocoducks1 &nd0 second0 (#(ll elieve in evolution %hen # see a monkey !ive irth to a
human ay1( This last one makes the same mistake as all the others0 plus the additional one of
thinkin! that maHor evolutionary chan!e happens overni!ht1
&s it happens0 t%o of these fallacies crop up next door to each other in the lon! list of
comments that follo% an article in the S"ndayTimes 7-ondon8 aout a television documentary on
'ar%in that # presentedA
'a%kins opinion on reli!ion is asurd since Evolution is nothin! more than a reli!ion itself
: you have to elieve %e all came from a sin!le cell 1 1 1 and that a snail can ecome a monkey etc1
)a )a : that(s the most lau!hale reli!ion yetKK
-oyce* Warwickshire* 9/
'a%kins should explain %hy science has failed to find the missin! links1 ,aith in unfounded
science is more fairy tale stuff than faith in God1
Bob* (as 1e$as* 9SA
This chapter %ill deal %ith all these related fallacies0 e!innin! %ith the silliest of all0 since
the ans%er to it %ill serve as an introduction to the others1
(Why doesn(t the fossil record contain a fronkeyB( Well0 of course0 monkeys are not
descended from fro!s1 *o sane evolutionist ever said they %ere0 or that ducks are descended from
crocodiles or vice versa1 "onkeys and fro!s share an ancestor0 %hich certainly looked nothin! like
a fro! and nothin! like a monkey1 "aye it looked a it like a salamander0 and %e do indeed have
salamander:like fossils datin! from the ri!ht time1 But that is not the point1 Every one of the
millions of species of animals shares an ancestor %ith every other one1 #f your understandin! of
evolution is so %arped that you think %e should expect to see a fronkey and a crocoduck0 you
should also %ax sarcastic aout the asence of a do!!ypotamus and an elephanFee1 #ndeed0 %hy
limit yourself to mammalsB Why not a kan!aroach 7intermediate et%een kan!aroo and cockroach80
or an octopard 7intermediate et%een octopus and leopard8B There(s an infinite numer of animal
names you can strin! to!ether in that %ay1 L Of course hippopotamuses are not descended from
do!s0 or vice versa1 ChimpanFees are not descended from elephants or vice versa0 Hust as monkeys
are not descended from fro!s1 *o modern species is descended from any other modern species 7if
%e leave out very recent splits81 .ust as you can find fossils that approximate to the common
ancestor of a fro! and a monkey0 so you can find fossils that approximate to the common ancestor
of elephants and chimpanFees1 )ere is one called Eomaia0 %hich lived in the early Cretaceous
period0 a little more than 255 million years a!o1
&s you can see0 Eomaia %as nothin! like a chimpanFee and nothin! like an elephant1
+a!uely like a shre%0 it proaly %as pretty similar to their common ancestor0 %ith %hich it %as
rou!hly contemporary0 and you can see that a lot of evolutionary chan!e has taken place alon! oth
path%ays from an Eomaia0 like ancestor to an elephant descendant0 and from the same Eomaia0 like
ancestor to a chimpanFee descendant1 But it is not in any sense an elephanFee1 #f it %ere0 it %ould
also have to e a do!atee0 for %hatever is the common ancestor of a chimpanFee and an elephant is
also the common ancestor of a do! and a manatee1 &nd it %ould also have to e an aardvapotamus0
for the same ancestor is also the common ancestor of an aardvark and a hippopotamus1 The very
idea of a do!atee 7or an elephanFee0 or an aardvapotamus or a kan!aroceros or a uffalion8 is deeply
unevolutionary and ridiculous1 So is a fronkey0 and it is a dis!race that the perpetrator of that little
%itlessism0 the &ustralian itinerant preacher .ohn "ackay0 has een tourin! British schools in 355;
and 35590 masEueradin! as a (!eolo!ist(0 teachin! innocent children that if evolution %ere true the
fossil record should contain (fronkeys(1
&n eEually ludicrous example is to e found in the "uslim apolo!ist )arun 6ahya(s
enormous0 lavishly produced0 !lossily illustrated and fatuously i!norant ook Atlas of #reation1
This ook oviously cost a fortune to produce0 %hich makes it all the more astoundin! that it %as
distriuted free to tens of thousands of science teachers0 includin! me1 *ot%ithstandin! the
prodi!ious sums of money spent on this ook0 the errors in it have ecome le!endary1 #n the service
of illustratin! the falsehood that most ancient fossils are indistin!uishale from their modern
counterparts0 6ahya sho%s a sea snake as an (eel( 7t%o animals so different that they are placed in
different classes of verterates80 a starfish as a (rittlestar( 7actually different classes of
echinoderms80 a saellid 7annelid8 %orm as a crinoid (sea lily( 7an echinodermA this pair come not
Hust from different phyla ut from different su:kin!doms0 so that they could hardly e more distant
from each other if they tried0 %hile still oth ein! animals8 and : est of all : a fishin! lure as a
(caddis fly( 7see colour pa!e ;81
But in addition to these !ems of partisan risiility0 the ook has a section on missin! links1
One picture is seriously offered to illustrate the fact that there is no intermediate form et%een a
fish and a starfish1 # find it impossile to elieve that the author seriously thinks evolutionists %ould
expect to find a transition et%een t%o such differin! animals as a starfish and a fish1 # therefore
cannot help suspectin! that he kno%s his audience all too %ell0 and is delierately and cynically
exploitin! their i!norance1
(#(-- BE-#E+E #* E+O-$T#O* W)E* & "O*DE6 G#+ES B#RT) TO & )$"&* B&B6(
Once a!ain0 humans are not descended from monkeys1 We share a common ancestor %ith
monkeys1 &s it happens0 the common ancestor %ould have looked a lot more like a monkey than a
man0 and %e %ould indeed proaly have called it a monkey if %e had met it0 some 3= million
years a!o1 But even thou!h humans evolved from an ancestor that %e could sensily call a monkey0
no animal !ives irth to an instant ne% species0 or at least not one as different from itself as a man is
from a monkey0 or even from a chimpanFee1 That isn(t %hat evolution is aout1 Evolution not only
is a !radual process as a matter of factJ it has to e !radual if it is to do any explanatory %ork1 )u!e
leaps in a sin!le !eneration : %hich is %hat a monkey !ivin! irth to a human %ould e : are almost
as unlikely as divine creation0 and are ruled out for the same reasonA too statistically improale1 #t
%ould e so nice if those %ho oppose evolution %ould take a tiny it of troule to learn the merest
rudiments of %hat it is that they are opposin!1
T)E PER*#C#O$S -EG&C6 O, T)E GRE&T C)&#* O, BE#*G
$nderlyin! much of the fallacious demand for (missin! links( is a medieval myth0 %hich
occupied men(s minds ri!ht up to the a!e of 'ar%in and stuornly confused them after it1 This is
the myth of the Great Chain of Bein!0 accordin! to %hich everythin! in the universe sat on a ladder0
%ith God at the top0 then archan!els0 then various ranks of an!els0 then human ein!s0 then animals0
then plants0 then do%n to stones and other inanimate creations1 Given that this !oes %ay ack to a
time %hen racism %as second nature0 # hardly need add that human ein!s %ere not all sittin! on
the same run!1 Oh no1 &nd of course males %ere a healthy run! aove females of their kind 7%hich
%as %hy # let myself !et a%ay %ith (occupied men's minds( in the openin! sentence of this section81
But it %as the alle!ed hierarchy %ithin the animal kin!dom that had the !reatest capacity to muddy
the %aters %hen the idea of evolution urst upon the scene1 #t seemed natural to suppose that (lo%er(
animals evolved into (hi!her( animals1 &nd if this %ere so0 %e should expect to see (links( et%een
them0 all the %ay up and do%n the (ladder(1 & ladder %ith lots of missin! run!s lacks conviction1 #t
is this ima!e of the run!less ladder that lurks ehind much of the scepticism aout (missin! links(1
But the entire ladder myth is deeply misconceived and un:evolutionary0 as # shall no% sho%1
So !lily do the phrases (hi!her animals( and (lo%er animals( trip off our ton!ues that it
comes as a shock to realiFe that0 far from effortlessly slottin! into evolutionary thinkin! as one
mi!ht suppose0 they %ere : and are : deeply antithetical to it1 We think %e kno% that chimpanFees
are hi!her animals and earth%orms are lo%er0 %e think %e(ve al%ays kno%n %hat that means0 and
%e think evolution makes it even clearer1 But it doesn(t1 #t is y no means clear that it means
anythin! at all1 Or if it means anythin!0 it means so many different thin!s as to e misleadin!0 even
)ere is a list of the more or less distinctly confusin! thin!s you mi!ht mean %hen you say0
for example0 that a monkey is (hi!her( than an earth%orm1
2 '%onkeys eoled from earthworms4' This is false0 Hust as it is false that humans evolved
from chimpanFees1 "onkeys and earth%orms share a common ancestor1
3 'The common ancestor of monkeys and earthworms was more like an earthworm than like
a monkey4' Well0 that potentially makes more sense1 6ou can even use the %ord (primitive( in a
semiprecise %ay0 if you define it as (resemlin! ancestors(0 and it is oviously true that some
modern animals are more primitive in this sense than others1 What that exactly means0 if you think
aout it0 is that the more primitive of a pair of species has chan!ed less since the common ancestor 7
all species0 %ithout exception0 share a common ancestor if you !o ack far enou!h81 #f neither
species has chan!ed dramatically more than the other0 the %ord (primitive( should not e used in
comparin! them1
#t(s %orth pausin! here to make a related point1 #t is hard to measure de!rees of resemlance1
&nd there is in any case no necessary reason %hy the common ancestor of t%o modern animals
should e more like one than the other1 #f you take t%o animals0 say a herrin! and a sEuid0 it is
possible that one of them resemles the common ancestor more than the other0 ut it doesn(t follo%
that this has to e the case1 There has een an exactly eEual amount of time for oth to have
diver!ed from the ancestor0 so the prior expectation of an evolutionist mi!ht e0 if anythin!0 that no
modern animal should e more primitive than any other1 We mi!ht expect oth of them to have
chan!ed to the same extent0 ut in different directions0 since the time of the shared ancestor1 This
expectation0 as it happens0 is often violated 7as in the case of monkey and earth%orm80 ut there is
no necessary reason %hy %e should expect it to e1 "oreover0 the different parts of animals don(t all
have to evolve at the same rate1 &n animal mi!ht e primitive from the %aist do%n ut hi!hly
evolved from the %aist up1 -ess facetiously0 one of them mi!ht e more primitive in its nervous
system0 the other more primitive in its skeleton1 *otice especially that (primitive( in the sense of
(resemlin! ancestors( does not have to !o %ith (simple( 7meanin! less complex81 & horse(s foot is
simpler than a human foot 7it has only a sin!le di!it instead of five0 for example80 ut the human
foot is more primitive 7the ancestor that %e share %ith horses had five di!its0 as %e do0 so the horse
has chan!ed more81 This leads us on to the next item in our list1
4 '%onkeys are cleerer :or prettier* hae lar$er $enomes* more complicated body plans*
etc4 etc4; than earthworms4' This kind of Foolo!ical snoery is a mess %hen you start tryin! to
apply it scientifically1 # mention it only ecause it is so readily confused %ith the other meanin!s0
and the est %ay to sort out confusion is to expose it1 6ou could ima!ine a lar!e numer of scales
alon! %hich you mi!ht rank animals : not Hust the four scales # have mentioned1 &nimals that are
hi!h on one of these ladders may or may not e hi!h on another1 "ammals certainly have lar!er
rains than salamanders0 ut they have smaller !enomes than some salamanders1
> '%onkeys are more like h"mans than earthworms are4' This is undeniale for the particular
example of monkeys and earth%orms1 But so %hatB Why should %e choose humans as the standard
a!ainst %hich %e Hud!e other or!anismsB &n indi!nant leech mi!ht point out that earth%orms have
the !reat virtue of ein! more like leeches than humans are1 'espite the Great Chain of Bein!(s
traditional rankin! of humans et%een animals and an!els0 there is no evolutionary Hustification for
the common assumption that evolution is someho% (aimed( at humans0 or that humans are
(evolution(s last %ord(1 #t is remarkale ho% commonly this vain!lorious assumption thrusts itself
for%ard1 &t its crudest level0 you meet it in the uiEuitously Euerulous0 (#f chimps evolved into us0
ho% come there are still chimps aroundB( #(ve already mentioned this0 and #(m not Hokin!1 # meet this
Euestion a!ain and a!ain and a!ain0 sometimes from apparently %ell:educated people1 L
= '%onkeys :and other 'hi$her' animals; are better at s"riin$ than earthworms :and other
'lower' animals;4' This doesn(t even e!in to e sensile0 or even true1 &ll livin! species have
survived at least into the present1 Some monkeys0 such as the exEuisite !olden tamarin0 are in
dan!er of !oin! extinct1 They are much less !ood at survivin! than earth%orms are1 Rats and
cockroaches flourish0 despite ein! re!arded y many people as (lo%er( than !orillas and oran!:
utans0 %hich are perilously close to extinction1
# hope #(ve said enou!h to sho% %hat nonsense it is to rank modern species on a ladder0 as
thou!h it %ere ovious %hat you meant y (hi!her( and (lo%er(0 and to sho% ho% thorou!hly
unevolutionary it is1 6ou can ima!ine lots and lots of laddersJ it mi!ht sometimes e sensile to
rank animals on at least some of the ladders separately0 ut the ladders are not %ell correlated %ith
each other0 and none of them has any ri!ht to e called an (evolutionary scale(1 We have seen the
historical temptation to crude errors such as (Why aren(t there any fronkeysB( But the pernicious
le!acy of the Great Chain of Bein! also feeds the challen!e (Where are the intermediates et%een
maHor animal !roupsB( and0 nearly as discreditaly0 underlies the tendency of evolutionists to
answer such a challen!e y trottin! out particular fossils0 such as Archaeopteryx0 the celerated
(intermediate et%een reptiles and irds(1 *evertheless0 there is somethin! else !oin! on underneath
the Archaeopteryx fallacy0 and it is of !eneral importanceJ so # shall !ive it a couple of para!raphs0
usin! Archaeopteryx as a particular example of a !eneral case1
Ioolo!ists have traditionally divided the verterates into classesA maHor divisions %ith
names like mammals0 irds0 reptiles and amphiians1 Some Foolo!ists0 called (cladists(0L insist that a
proper class must consist of animals all of %hom share a common ancestor %hich elon!ed to that
class and %hich has no descendants outside that !roup1 The irds %ould e an example of a !ood
&ll irds are descended from a sin!le ancestor that %ould also have een called a ird and
%ould have shared %ith modern irds the key dia!nostic characters : feathers0 %in!s0 a eak0 etc1
The animals commonly called reptiles are not a !ood class in this sense1 This is ecause0 at least in
conventional taxonomies0 the cate!ory explicitly excl"des irds 7they constitute their o%n class8 and
yet some (reptiles( as conventionally reco!niFed 7e1!1 crocodiles and dinosaurs8 are closer cousins to
irds than they are to other (reptiles( 7e1!1 liFards and turtles81 #ndeed0 some dinosaurs are closer
cousins to irds than they are to other dinosaurs1 (Reptiles(0 then0 is an artificial class0 ecause irds
are artificially excl"ded1 #n a strict sense0 if %e %ere to make reptiles a truly natural class0 %e should
have to include irds as reptiles1 Cladistically inclined Foolo!ists avoid the %ord (reptiles(
alto!ether0 splittin! them into &rchosaurs 7crocodiles0 dinosaurs and irds80 -epidosaurs 7snakes0
liFards and the rare Sphenodon of *e% Iealand8 and Testudines 7turtles and tortoises81 *on:
cladistically inclined Foolo!ists are happy to use a %ord like (reptile( ecause they find it
descriptively useful0 even if it does artificially exclude the irds1
But %hat is it aout the irds that tempts us to hive them off from the reptilesB What is it
that seems to Hustify esto%in! on irds the accolade of (class(0 %hen they are0 evolutionarily
speakin!0 Hust one ranch %ithin reptilesB #t is the fact that the immediately surroundin! reptiles0
irds( close nei!hours on the tree of life0 happen to e extinct0 %hile the irds0 alone of their kind0
marched on1 The closest relatives of irds are all to e found amon! the lon!extinct dinosaurs1 #f a
%ide variety of dinosaur linea!es had survived0 irds %ould not stand outA they %ould not have een
elevated to the status of their o%n class of verterates0 and %e %ould not e askin! any such
Euestion as (Where are the missin! links et%een reptiles and irdsB( Archaeopteryx %ould still e a
nice fossil to have in your museum0 ut it %ould not play its present starrin! role as the stock
ans%er to 7%hat %e can no% see is8 an empty challen!eA (Produce your intermediates1( #f the cards
of extinction had fallen differently0 there %ould Hust e lots of dinosaurs runnin! aout0 includin!
some feathered0 flyin!0 eaked dinosaurs called irds1 &nd indeed0 fossiliFed feathered dinosaurs
are no% increasin!ly ein! discovered0 so it is ecomin! vividly clear that there really is no maHor
(Produce your missin! linkK( challen!e to %hich Archaeopteryx is the reply1
-et(s proceed0 no%0 to some of the maHor transitions in evolution0 %here (links( have een
alle!ed to e (missin!(1
$P ,RO" T)E SE&
Short of rocketin! into space0 it is hard to ima!ine a older or more life:chan!in! step than
leavin! the %ater for dry land1 The t%o life:Fones are different in so many %ays that movin! from
one to the other demands a radical shift in almost all parts of the ody1 Gills that are !ood at
extractin! oxy!en from %ater are all ut useless in air0 and lun!s are useless in %ater1 "ethods of
propulsion that are speedy0 !raceful and efficient in %ater are dan!erously clumsy on land0 and vice
versa1 *o %onder (fish out of %ater( and (like a dro%nin! man( have oth ecome proverial
phrases1 &nd no %onder (missin! links( in this re!ion of the fossil record are of more than ordinary
#f you !o ack far enou!h0 everythin! lived in the sea : %atery0 salty alma mater of all life1
&t various points in evolutionary history0 enterprisin! individuals from many different animal
!roups moved out on to the land0 sometimes eventually to the most parched deserts0 takin! their
o%n private sea %ater %ith them in lood and cellular fluids1 #n addition to the reptiles0 irds0
mammals and insects %e see all around us0 other !roups that have succeeded in makin! the !reat
trek out of life(s %atery %om include scorpions0 snails0 crustaceans such as %oodlice and land
cras0 millipedes and centipedes0 spiders and their kin0 and at least three phyla of %orms1 &nd %e
mustn(t for!et the plants0 onlie e!etters of usale caron0 %ithout %hose prior invasion of the land
none of the other mi!rations could have happened1
,ortunately the transitional sta!es of our exodus0 as fish emer!ed on to the land0 are
eautifully documented in the fossil record1 So are the transitional sta!es !oin! the other %ay much
later0 as the ancestors of %hales and du!on!s forsook their hard:%on home on dry land and returned
to their ancestral seas1 #n oth cases0 links that once %ere missin! no% aound and !race our
When %e say that (fish( emer!ed on to the land0 %e have to rememer that (fish(0 like
(reptiles(0 do not constitute a natural !roup1 ,ish are defined y exclusion1 ,ish are all the verterates
except those that moved on to the land1 Because all the early evolution of verterates took place in
%ater0 it is not surprisin! that most of the survivin! ranches of the verterate tree are still in the
sea1 &nd %e still call them (fish( even %hen they are only distantly related to other (fish(1 Trout and
tuna are closer cousins to humans than they are to sharks0 ut %e call them all (fish(1 &nd lun!fish
and coelacanths are closer cousins to humans than they are to trout and tuna 7and of course sharks8
ut0 a!ain0 %e call them (fish(1 Even sharks are closer cousins to humans than they are to lampreys
and ha!fish 7the only modern survivors of the once thrivin! and diverse !roup of Ha%less fishes8 ut
a!ain0 %e call them all fish1 +erterates %hose ancestors never ventured on to land all look like
(fish(0 they all s%im like fish 7unlike dolphins0 %hich s%im %ith an up:and:do%n endin! of the
spine instead of side to side like a fish80 and they all0 # suspect0 taste like fish1
To an evolutionist0 as %e Hust sa% in the example of reptiles and irds0 a (natural( !roup of
animals is a !roup all of %hose memers are closer cousins to each other than they are to all non:
memers of the !roup1 (Birds(0 as %e sa%0 are a natural !roup0 since they share a most recent
common ancestor that is not shared y any non:ird1 By the same definition0 (fish( and (reptiles( are
not natural !roups1 The most recent common ancestor of all (fish( is shared y many non:fish too1 #f
%e push our distant cousins the sharks to one side0 %e mammals elon! to a natural !roup that
includes all modern ony fish 7ony as opposed to cartila!inous sharks81 #f %e then push to one side
the ony (ray:finned fishes( 7salmon0 trout0 tuna0 an!el fishA Hust aout all the fish you are likely to
see that are not sharks80 the natural !roup to %hich %e elon! includes all land verterates plus the
so:called loe:finned fishes1 #t is from the ranks of the loe:finned fishes that %e spran!0 and %e
must no% pay special attention to the loefins1
-oefins today have d%indled to the lun!fishes and the coelacanths 7(d%indled( as (fish(0 that
is0 ut mi!htily expanded on landA %e land verterates are aerrant lun!fish81 They are (loefins(
ecause their fins are like le!s rather than the ray fins of familiar fishes1 #ndeed0 !ld ,o"rle$s %as
the title of a popular ook on coelacanths %ritten y .1 -1 B1 Smith0 the South &frican iolo!ist
most responsile for rin!in! them to the %orld(s attention after the first live one %as dramatically
discovered in 294; in the catch of a South &frican tra%lerA (# %ould not have een more surprised if
# had seen a dinosaur %alkin! do%n the street1( Coelacanths had een kno%n efore0 as fossils0 ut
they had een thou!ht extinct since the time of the dinosaurs1 Smith movin!ly %rote of the moment
%hen he first cast eyes on this astonishin! find0 to %hich he had een summoned y its discoverer0
"ar!aret -atimer 7he later named it (atimeria80 to !ive his expert opinionA
We %ent strai!ht to the "useum1 "iss -atimer %as out for the moment0 the caretaker
ushered us into the inner room and there %as the : Coelacanth0 yes0 GodK &lthou!h # had come
prepared0 that first si!ht hit me like a %hite:hot last and made me feel shaky and Eueer0 my ody
tin!led1 # stood as if stricken to stone1 6es0 there %as not a shado% of dout0 scale y scale0 one y
one0 fin y fin0 it %as a true Coelacanth1 #t could have een one of those creatures of 355 million
years a!o come alive a!ain1 # for!ot everythin! else and Hust looked and looked0 and then almost
fearfully %ent close up and touched and stroked0 %hile my %ife %atched in silence1 "iss -atimer
came in and !reeted us %armly1 #t %as only then that speech came ack0 the exact %ords # have
for!otten0 ut it %as to tell them that it %as true0 it %as really true0 it %as unEuestionaly a
Coelacanth1 *ot even # could dout any more1
Coelacanths are closer cousins to us than they are to most fish1 They have chan!ed
some%hat since the time of our shared ancestor0 ut not enou!h to e moved out of the cate!ory of
animals that0 colloEuially and to a fisherman0 %ould e classified as fish1 But they0 and lun!fish0 are
definitely closer cousins to us than to trout0 tuna and the maHority of fish1 Coelacanths and lun!fish
are examples of (livin! fossils(1
*evertheless0 %e are not descended from lun!fish0 or from coelacanths1 We share an
ancestor %ith lun!fish0 %hich looked more like a lun!fish than it looked like us1 But it didn(t look
much like either1 -un!fish may e livin! fossils0 ut they are still not very like our ancestors1 #n the
Euest for those0 %e must instead seek real fossils in the rocks1 &nd in particular %e are interested in
fossils from the 'evonian era that capture the transition et%een %ater:d%ellin! fish and the first
verterates to live on land1 Even amon! real fossils0 %e %ould e too optimistic if %e hoped
literally to find our ancestors1 We can0 ho%ever0 hope to find cousins of our ancestors that are
sufficiently close to tell us approximately %hat they %ere like1
One of the most famous !aps in the fossil record : conspicuous enou!h to have een !iven a
name0 (Romer(s Gap( 7&1 S1 Romer %as a famous &merican palaeontolo!ist80 stretches from aout
4<5 million years a!o0 at the end of the 'evonian period0 to aout 4>5 million years a!o0 in the
early part of the Caroniferous0 the (Coal "easures(1 &fter Romer(s Gap0 %e find uneEuivocal
amphiians cra%lin! throu!h the s%amps0 a rich radiation of salamander:like animals0 some of
them as lar!e as crocodiles0 %hich they superficially resemled1 #t seems to have een an a!e of
!iants0 for there %ere dra!onflies %ith a %in! span as lon! as my arm0 the lar!est insects that ever
lived1L Startin! aout 4>5 million years a!o0 %e mi!ht almost call the Caroniferous the amphiian
eEuivalent of the a!e of dinosaurs1 Before that0 ho%ever0 %as Romer(s Gap1 &nd efore his !ap0
Romer could see only fish0 loe:finned fish0 livin! in %ater1 Where %ere the intermediates0 and
%hat led them to venture out on to the landB
"y under!raduate ima!ination at Oxford %as fired y the lectures of the prodi!iously
kno%led!eale )arold Pusey %ho0 despite his dry and prolon!ed delivery0 had a !ift for seein!
eyond dry ones to the flesh:and:lood animals that had to make a livin! in some departed %orld1L
)is evocation of %hat drove some loe:finned fish to develop lun!s and le!s0 %hich %as derived
from Romer himself0 made memorale sense to my student ears0 and it still makes sense to me even
thou!h it is less fashionale amon! modern palaeontolo!ists than it %as in Romer(s time1 Romer0
and Pusey0 envisa!ed annual drou!hts durin! %hich lakes and ponds and streams dried up0 only to
flood a!ain the follo%in! year1 ,ishes that made their livin! in %ater could enefit from a
temporary aility to survive on land0 %hile they dra!!ed themselves from a shallo% lake or pond
that %as threatened %ith imminent desiccation to a deeper one in %hich they could survive until the
next %et season1 On this vie%0 our ancestors didn(t so much emer!e on to the dry land as use the dry
land as a temporary rid!e to escape ack into the %ater1 "any modern animals do the same1
Rather unfortunately0 Romer introduced his theory %ith a preamle %hose purpose %as to
sho% that the 'evonian era %as a time of drou!ht1 ConseEuently0 %hen more recent evidence
undermined this assumption0 it seemed to undermine the %hole Romer theory1 )e(d have done
etter to omit the preamle0 %hich %as0 in any case0 overkill1 &s # ar!ued in The Ancestor's Tale0
the theory still %orks0 even if the 'evonian %as less drou!ht:ridden than Romer ori!inally thou!ht1
-et us0 in any case0 return to the fossils themselves1 They trickle sparsely throu!h the late
'evonian0 the period immediately precedin! the CaroniferousA tantaliFin! traces of (missin! links(0
animals that %ent some %ay to%ards rid!in! the !ap et%een the loe:finned fishes that %ere so
aundant in 'evonian seas0 and the amphiians that later slithered throu!h the Caroniferous
s%amps1 On the fish side of the !ap0 E"sthenopteron %as discovered in 2;;2 in a collection of
fossils from Canada1 #t seems to have een a surface:huntin! fish and proaly didn(t ever come on
land0 not%ithstandin! some early ima!inative reconstructions1 *evertheless0 it did have several
anatomical similarities to the amphiians of =5 million years later0 includin! its skull ones0 its
teeth and0 aove all0 its fins1 &lthou!h they %ere proaly used for s%immin! and not %alkin!0 the
ones follo%ed the typical pattern of a tetrapod 7the name !iven to all land verterates81 #n the
forelim0 a sin!le humerus %as Hoined to t%o ones0 the radius and ulna0 Hoined to lots of little
ones0 %hich %e tetrapods %ould call carpals0 metacarpals and fin!ers1 &nd the hind lim sho%s a
similar tetrapod:like pattern1
Then0 near the amphiian side of the !ap0 some 35 million years later0 at the order et%een
the 'evonian and Caroniferous0 !reat excitement %as caused y the 2943 discovery in Greenland
of &chthyoste$a4 'on(t e misled y thou!hts of cold and ice0 y the %ay1 Greenland in the days of
&chthyoste$a %as on the eEuator1 &chthyoste$a %as first reconstructed y the S%edish
palaeontolo!ist Erik .arvik in 29==0 and a!ain he portrayed it as closer to a land:d%eller than
modern experts do1 The most recent reconstruction0 y Per &hler! at .arvik(s old university of
$ppsala0 places &chthyoste$a mostly in the %ater0 althou!h it proaly made occasional forays on to
the land1 *evertheless0 it looked more like a !iant salamander than a fish0 and it had the flat head
that is so characteristic of amphiians1 $nlike all modern tetrapods0 %hich have five fin!ers and
toes 7at least in the emryo0 althou!h they may lose some in the adult80 &chthyoste$a had seven toes1
#t seems that the early tetrapods enHoyed more freedom to (experiment( %ith varyin! numers of
di!its than %e have today1 Presumaly at some point the emryolo!ical processes fixed upon five0
and a step %as taken that %as hard to reverse1 &lthou!h0 admittedly0 not as hard as all that1 There
are individual cats0 and indeed humans0 %ho have six toes1 These extra toes proaly arise throu!h
a duplication error in emryolo!y1
&nother excitin! discovery0 also from tropical Greenland and also datin! from the oundary
et%een the 'evonian and the Caroniferous0 %as Acanthoste$a4 Acanthoste$a0 too0 had a flat0
amphiian skull and tetrapod:like limsJ ut it too departed0 and even further than &chthyoste$a0
from %hat %e no% think of as the fivefin!er standard1 #t had ei!ht di!its1 The scientists most
responsile for our kno%led!e of it0 .enny Clack and "ichael Coates of Camrid!e $niversity0
elieve that0 like &chthyoste$a0 Acanthoste$a %as lar!ely a %ater:d%eller0 ut it had lun!s and its
lims stron!ly su!!est that it could cope %ith land as %ell as %ater if it had to1 &!ain0 it looked
pretty much like a !iant salamander1 "ovin! ack no% to the fish side of the divide0 Panderichthys*
also from the late 'evonian0 is also sli!htly more amphiian:like0 and sli!htly less fish:like0 than
E"sthenopteron1 But if you sa% it you %ould surely %ant to call it a fish rather than a salamander1
So0 %e are left %ith a !ap et%een Panderichthys0 the amphiian:like fish0 and
Acanthoste$a0 the fish:like amphiian1 Where is the (missin! link( et%een themB & team of
scientists from the $niversity of Pennsylvania0 includin! *eil Shuin and Ed%ard 'aeschler0 set
out to find it1 Shuin made their Euest the asis for a deli!htful series of reflections on human
evolution in his ook <o"r &nner ,ish1 They delierately thou!ht aout %here mi!ht e the est
place to look0 and carefully chose a rocky area of exactly the ri!ht late 'evonian a!e in the
Canadian &rctic1 There they %ent : and struck Foolo!ical !old1 TiktaalikK & name never to e
for!otten1 #t comes from an #nuit %ord for a lar!e fresh%ater fish1 &s for the specific name0 roseae0
let me tell a cautionary tale a!ainst myself1 When # first heard the name0 and sa% photo!raphs like
the one reproduced on colour pa!e 250 my mind immediately leapt to the 'evonian0 the (Old Red
Sandstone(0 the colour of the eponymous county of 'evon0 the colour of Petra 7(& rose:red city0 half
as old as time(81 &las0 # %as Euite %ron!1 The photo!raph exa!!erates the rosy !lo%1 The name %as
chosen in honour of a enefactor %ho helped finance the expedition to the &rctic 'evonian1 # %as
privile!ed to e sho%n Tiktaalikroseae y 'r 'aeschler %hen # had lunch %ith him in Philadelphia0
shortly after its discovery0 and the lifelon! Foolo!ist in me : or perhaps my inner fish : %as moved
to speechlessness1 Throu!h rose:tinted spectacles # ima!ined # %as !aFin! upon the face of my
direct ancestor1 $nrealistic as that %as0 this not:so:rose:red fossil %as proaly as close as # %as
!oin! to !et to meetin! a real dead ancestor half as old as time1
#f you %ere to meet a real live Tiktaalik0 snout to snout0 you mi!ht start ack as if threatened
y a crocodile0 for that is %hat its face resemled1 & crocodile(s head on a salamander(s trunk0
attached to a fish(s rear end and tail1 $nlike any fish0 Tiktaalik had a neck1 #t could turn its head1 #n
almost every particular0 Tiktaalik is the perfect missin! link : perfect0 ecause it almost exactly
splits the difference et%een fish and amphiian0 and perfect ecause it is missin! no lon!er1 We
have the fossil1 6ou can see it0 touch it0 try to appreciate the a!e of it : and fail1
# "$ST GO 'OW* TO T)E SE& &G&#*L
The move from %ater to land launched a maHor redesi!n of every aspect of life0 from
reathin! to reproductionA it %as a !reat trek throu!h iolo!ical space1 *evertheless0 %ith %hat
seems almost %anton perversity0 a !ood numer of thorou!h!oin! land animals later turned around0
aandoned their hard:earned terrestrial retoolin!0 and trooped ack into the %ater a!ain1 Seals and
sea lions have only !one part:%ay ack1 They sho% us %hat the intermediates mi!ht have een like0
on the %ay to extreme cases such as %hales and du!on!s1 Whales 7includin! the small %hales %e
call dolphins80 and du!on!s %ith their close cousins the manatees0 ceased to e land creatures
alto!ether and reverted to the full marine haits of their remote ancestors1 They don(t even come
ashore to reed1 They do0 ho%ever0 still reathe air0 havin! never developed anythin! eEuivalent to
the !ills of their earlier marine pro!enitors1 Other animals that have returned from land to %ater0 at
least some of the time0 are pond snails0 %ater spiders0 %ater eetles0 crocodiles0 otters0 sea snakes0
%ater shre%s0 Galapa!os fli!htless cormorants0 Galapa!os marine i!uanas0 yapoks 7aEuatic
marsupials from South &merica80 platypuses0 pen!uins and turtles1
Whales %ere lon! an eni!ma0 ut recently our kno%led!e of %hale evolution has ecome
rather rich1 "olecular !enetic evidence 7see Chapter 25 for the nature of this kind of evidence8
sho%s that the closest livin! cousins of %hales are hippos0 then pi!s0 then ruminants1 Even more
surprisin!ly0 the molecular evidence sho%s that hippos are more closely related to %hales than they
are to the cloven:hoofed animals 7such as pi!s and ruminants8 %hich look much more like them1
This is another example of the mismatch that can sometimes arise et%een closeness of cousinship
and de!ree of physical resemlance1 We noted it aove in connection %ith fish that are closer
cousins to us than they are to other fish1 #n that case0 the anomaly arose ecause our linea!e left the
%ater for the land0 and conseEuently sur!ed a%ay in evolution0 leavin! our close fish cousins0 the
lun!fish and coelacanths0 resemlin! our more distant fish cousins ecause they all stayed in the
%ater1 *o% %e meet the same phenomenon a!ain0 ut in reverse1 )ippos stayed0 at least partly0 on
land0 and so still resemle their more distant land:d%ellin! cousins0 the ruminants0 %hile their
closer cousins0 the %hales0 took off into the sea and chan!ed so drastically that their affinities %ith
hippos escaped all iolo!ists except molecular !eneticists1 &s %hen their remote fishy ancestors
ori!inally %ent in the other direction0 it %as a it like takin! off into space0 or at least like launchin!
a alloon0 as the ancestors of %hales floated free of the constrainin! urden of !ravity and severed
their moorin!s to dry land1
&t the same time0 the once rather scanty fossil record of %hale evolution has een
convincin!ly filled out0 mostly y a ne% trove from Pakistan1 )o%ever0 the story of fossil %hales
has een so %ell treated in other recent ooks0 for example 'onald Prothero(s Eol"tion= What the
,ossils Say and Why it %atters0 and0 more recently0 .erry Coyne(s Why Eol"tion is Tr"e0 that #
have decided not to cover the same details here1 #nstead0 # have confined myself to one dia!ram
7elo%80 taken from Prothero(s ook0 sho%in! a seEuence of fossils ordered in time1 *ote the
careful %ay the picture is dra%n1 #t is temptin! : and older ooks used to do this : to dra% seEuences
of fossils %ith arro%s from older to youn!er ones1 But noody can say0 for example0 that
Amb"locet"s %as descended from Pakicet"s1 Or that Basilosa"r"s %as descended from Rodhocet"s1
#nstead0 the dia!ram follo%s the more cautious policy of su!!estin! that0 for example0 %hales are
descended from a contemporary cousin of Amb"locet"s%hich %as proaly rather like Amb"locet"s
7and mi!ht even have een Amb"locet"s81 The fossils sho%n are representative of various sta!es of
%hale evolution1 The !radual disappearance of the hind lims0 the transformation of the front lims
from %alkin! le!s to s%immin! fins0 and the flattenin! of the tail into flukes0 are amon! the
chan!es that emer!ed in ele!ant cascade1
Fossil ,hales
Fi'(re 12.131 Evolution of %hales from land creatures0 sho%in! the numerous transitional
fossils no% documented from the Eocene eds of &frica and Pakistan1 7'ra%in! y Carl Buell8
That(s all #(m !oin! to say aout the fossil history of %hales0 ecause it has een so %ell
treated in the ooks # mentioned1 The other0 less numerous and diverse ut Hust as thorou!hly
aEuatic !roup of marine mammals0 the sirenians : du!on!s and manatees : are not so %ell
documented in the fossil record0 ut one outstandin!ly eautiful (missin! link( has recently een
discovered1 Rou!hly contemporary %ith Amb"locet"s0 the Eocene (%alkin! %hale(0 is Pe2osiren0 the
(%alkin! manatee( fossil from .amaica1 #t looks pretty much like a manatee or du!on!0 except that it
has proper %alkin! le!s oth front and rear0 %here they have flippers in the front and no lims at all
in the rear1 The picture opposite sho%s a modern du!on! skeleton aove0 Pe2osiren elo%1
.ust as %hales are related to hippos0 so sirenians are related to elephants0 as a !reat deal of
evidence0 includin! the all:important molecular evidence0 attests1 Pe2osiren0 ho%ever0 proaly
lived like a hippo0 spendin! much of its time in %ater and usin! its le!s to %alk on the ottom as
%ell as s%im1 The skull is unmistakaly sirenian1 Pe2osiren may or may not e the actual ancestor
of modern manatees and du!on!s0 ut it is certainly excellent castin! for the role1
This ook %as aout to !o to the printer %hen excitin! ne%s came in0 from the Hournal
.at"re0 of a ne% fossil from the Canadian &rctic0 plu!!in! a !ap in the ancestry of modern seals0
sea lions and %alruses 7collectively (pinnipeds(81 & sin!le skeleton0 aout <= per cent complete0
P"i6ila darwini dates from the early "iocene epoch 7aout 35 million years a!o81 That(s recent
enou!h that the map of the %orld %as almost the same as today1 So this early sealGsea lion 7they had
not diver!ed yet8 %as an &rctic animal0 a deniFen of cold %ater1 Evidence su!!ests that it lived and
fished in fresh %ater 7like most otters except the famous sea otters of California80 rather than in the
sea 7like most modern seals except the famous -ake Baikal seal81 P"i6ila did not have flippers0 ut
%eed feet1 #t proaly ran like a do! on land 7very unlike a modern pinniped8 ut spent much of
its time in %ater0 %here it s%am like a do!0 unlike either of the t%o styles adopted respectively y
modern seals and sea lions1 P"i6ila neatly straddles the !ap et%een land and %ater in the ancestry
of pinnipeds1 #t is yet another deli!htful addition to our !ro%in! list of (links( that are no lon!er
4o%ern %('on'
Pezosiren) anient %('on'
# no% %ant to turn to another !roup of animals that returned from the land to the %aterA a
particularly intri!uin! example ecause some of them later reversed the process and returned to the
land a second timeK Sea turtles are0 in one important respect0 less fully !iven ack to the %ater than
%hales or du!on!s0 for they still lay their e!!s on eaches1 -ike all verterate returners to the %ater0
turtles haven(t !iven up reathin! air0 ut in this department some of them !o one etter than
%hales1 These turtles extract additional oxy!en from the %ater throu!h a pair of chamers at their
rear end that are richly supplied %ith lood vessels1 One &ustralian river turtle0 indeed0 !ets the
maHority of its oxy!en y reathin! 7as an &ustralian %ould not hesitate to say8 throu!h its arse1
Before !oin! any further0 # can(t escape a tiresome point of terminolo!y0 and a re!rettale
vindication of Geor!e Bernard Sha%(s oservation that (En!land and &merica are t%o countries
divided y a common lan!ua!e1( #n Britain0 turtles live in the sea0 tortoises live on land and
terrapins live in fresh or rackish %ater1 #n &merica all these animals are (turtles(0 %hether they live
on land or in %ater1 (-and turtle( sounds odd to me0 ut not to an &merican0 for %hom tortoises are
the suset of turtles that live on land1 Some &mericans use (tortoise( in a strict taxonomic sense to
refer to the Testudinidae0 %hich is the scientific name for modern land tortoises1 #n Britain0 %e(d e
inclined to call any land:d%ellin! chelonian a tortoise0 %hether it is a memer of the Testudinidae
or not 7as %e shall see0 there are fossil (tortoises( that lived on land ut are not memers of the
Testudinidae81 #n %hat follo%s0 #(ll try to avoid confusion0 makin! allo%ance for readers in Britain
and &merica 7and &ustralia0 %here the usa!e is different a!ain80 ut it(s hard1 The terminolo!y is a
mess0 to put it mildly1 Ioolo!ists use (chelonians( for all these animals0 turtles0 tortoises and
terrapins0 %hichever version of En!lish %e speak1
The most instantly noticeale feature of chelonians is their shell1 )o% did it evolve0 and
%hat did the intermediates look likeB Where are the missin! linksB What 7a creationist Fealot mi!ht
ask8 is the use of half a shellB Well0 amaFin!ly0 a ne% fossil has Hust een descried0 %hich
eloEuently ans%ers that Euestion1 #t made its deut in the Hournal .at"re in the nick of time efore #
had to hand this ook over to the pulishers1 #t %as an aEuatic turtle0 found in late Triassic
sediments in China0 and its a!e is estimated at 335 million years1 #ts name is !dontochelys
semitestacea0 from %hich you may deduce that0 unlike a modern turtle or tortoise0 it had teeth0 and
it did indeed have half a shell1 #t also had a much lon!er tail than a modern turtle or tortoise1 &ll
three of these features mark it out as prime (missin! link( material1 The elly %as covered y a shell0
the so:called plastron0 in pretty much the same %ay as that of a modern sea turtle1 But it almost
completely lacked the dorsal portion of the shell0 kno%n as the carapace1 #ts ack %as presumaly
soft0 like a liFard(s0 althou!h there %ere some hard0 ony its alon! the middle aove the ackone0
as in a crocodile0 and the ris %ere flattened0 as thou!h (tryin!( to form the evolutionary e!innin!s
of a carapace1
&nd here %e have an interestin! controversy1 The authors of the paper that introduced
!dontochelys to the %orld0 -i0 Wu0 Rieppel0 Wan! and Ihao 7for revity0 #(ll call them the Chinese
authors0 althou!h Rieppel is not Chinese80 think that their animal %as indeed half%ay to%ards
acEuirin! a shell1 Others dispute !dontochelys(s claim to demonstrate that the shell evolved in
%ater1 .at"re has the admirale custom of commissionin! experts other than the authors to %rite a
commentary on the %eek(s more interestin! articles0 %hich they pulish in a section called (*e%s
and +ie%s(1 The (*e%s and +ie%s( commentary on the !dontochelys paper is y t%o Canadian
iolo!ists0 Roert ReisF and .ason )ead0 and they offer an alternative interpretation1 "aye the
%hole shell had already evolved on land0 efore !dontochelys(s ancestors %ent ack to the %ater1
&nd maye !dontochelys lost its carapace after returnin! to the %ater1 ReisF and )ead point out
that some of today(s sea turtles0 for example the !iant leatherack turtle0 have lost or !reatly reduced
the carapace0 so their theory is Euite plausile1
# need to di!ress for a rief aside on the Euestion0 (What is the use of half a shellB( #n
particular0 %hy %ould !dontochelys e armoured elo% ut not aoveB Perhaps ecause dan!er
threatened from elo%0 %hich %ould su!!est that these creatures spent a lot of their time s%immin!
near the surface : and of course they had to come to the surface to reathe0 any%ay1 Sharks today
often attack from elo%0 sharks %ould have een a menacin!ly important part of the %orld of
!dontochelys0 and there(s no reason to suppose that their huntin! haits %ere different in those
times1 &s a parallel example0 one of the most surprisin! achievements of evolution0 the extra pair of
eyes in the fish Bathylychnops 7see over80 is proaly aimed at detectin! predatory attacks from
elo%1 The main eye looks out%ards0 as in any ordinary fish1 But each of the t%o main eyes has an
extra little eye0 complete %ith lens and retina0 tucked into its lo%er side1 #f Bathylychnops can !o to
the troule 7you kno% %hat # mean0 don(t e pedantic8 of !ro%in! a %hole extra pair of eyes0
presumaly to look out for attacks comin! from elo%0 it seems Euite plausile that !dontochelys
mi!ht !ro% armour aimed at fendin! off attacks from the same direction1 The plastron makes sense1
&nd if you %ant to say0 yes0 ut %hy not have a carapace on top as %ell0 Hust to e extra safe0 the
reply is easy1 Shells are heavy and cumersome0 they are costly to !ro% and costly to carry around1
There are al%ays trade:offs in evolution1 ,or land tortoises0 the trade:off ends up favourin! stout0
heavy armour aove as %ell as elo%1 ,or many sea turtles0 the tradeoff favours a stron! plastron
underneath ut li!hter armour on top1 &nd it is a plausile su!!estion that !dontochelys Hust carried
that trend a it further1
athylychnops!e/tra eye
#f0 on the other hand0 the Chinese authors are ri!ht that !dontochelys %as on its %ay to
evolvin! a full shell0 and that the shell evolved in %ater0 it %ould seem to follo% that modern land
tortoises0 %hich have %ell:developed shells0 are descended from %ater turtles1 This0 as %e shall see0
is proaly true1 But it is remarkale0 ecause it means that today(s land tortoises represent a second
mi!ration from %ater to land1 *oody has ever claimed that any %hales0 or du!on!s0 ret"rned to the
land after invadin! the %ater1 The alternative story for land tortoises is that they %ere on land all
alon! and independently evolved the shell0 in parallel to their aEuatic cousins1 This is y no means
impossileJ ut0 as it happens0 %e have !ood reason to elieve that sea turtles did indeed return to
the land for a second !o at ecomin! land tortoises1
Fa"ily tree o$ tortoises an% t(rtles
*ol% R land
normal R aEuatic
#f you dra% out the family tree of all modern turtles and tortoises0 ased on molecular and
other comparisons0 nearly all the ranches are aEuatic 7normal type81 -and tortoises are represented
y old type0 and you can see that today(s land tortoises constitute a sin!le ranch0 the Testudinidae0
deeply nested %ithin rich ranchin!s of other%ise aEuatic chelonians1 &ll their close cousins are
aEuatic1 "odern land tortoises are a sin!le t%i! on the ush of other%ise aEuatic turtles1 Their
aEuatic ancestors turned turtle and trooped ack on to the land1 This fact is compatile %ith the
hypothesis that the shell evolved in %ater0 in a creature like !dontochelys1 But no% %e have
another difficulty1 #f you look at the family tree0 you(ll notice that0 in addition to the Testudinidae
7all modern land tortoises8 there are t%o fossil !enera of fully shelled animals called
Pro$anochelysL and Palaeochersis1 These are dra%n as land:d%ellers0 for reasons %e shall come to
in the next para!raph1 They lie ri!ht outside the ranches representin! the %ater turtles1 #t %ould
seem that these t%o !enera are anciently terrestrial1
Before !dontochelys %as discovered0 these t%o fossils %ere the oldest kno%n chelonians1
-ike !dontochelys they lived in the late Triassic0 ut aout 2= million years later than
!dontochelys1 Some authorities have reconstructed them as livin! in fresh %ater0 ut recent
evidence does indeed place them on land0 as indicated y old type on the dia!ram1 6ou mi!ht
%onder ho% %e tell %hether fossil animals0 especially if only fra!ments are found0 lived on land or
in %ater1 Sometimes it(s pretty ovious1 #chthyosaurs %ere reptilian contemporaries of the
dinosaurs0 %ith fins and streamlined odies1 The fossils look like dolphins and they surely lived like
dolphins0 in the %ater1 With turtles and tortoises it is a little less ovious1 &s you mi!ht expect0 the
i!!est !ivea%ay is their lims1 Paddles really are rather different from %alkin! le!s1 Walter .oyce
and .acEues Gauthier0 of 6ale $niversity0 took this common:sense intuition and provided the
numers to support it1 They took three key measurements in the arm and hand ones of seventy:one
species of livin! chelonians1 #(ll resist the temptation to explain their ele!ant calculations0 ut their
conclusion %as clear1 These animals had had %alkin! le!s0 not paddles1 #n British En!lish0 they
%ere (tortoises(0 not (turtles(1 They lived on land1 They %ere only distant cousins0 ho%ever0 of
modern land tortoises1
*o% %e seem to have a prolem1 #f0 as the authors of the paper descriin! !dontochelys
elieve0 their half:shelled fossil sho%s that the shell evolved in %ater0 ho% do %e explain t%o
!enera of fully shelled (tortoises( on land0 2= million years laterB $ntil the discovery of
!dontochelys0 # %ould not have hesitated to say that Pro$anochelys and Palaeochersis %ere
representative of the land:d%ellin! ancestral type before the return to %ater1 The shell evolved on
land1 Some shelled tortoises returned to the sea0 as seals0 %hales and du!on!s %ere later to do1
Others stayed on land0 ut %ent extinct1 &nd then some sea turtles returned to the land0 to !ive rise
to all modern land tortoises1 That(s %hat # %ould have said : indeed %hat # did say in the earlier
draft of this chapter that preceded the announcement of !dontochelys1 But !dontochelys thro%s
speculation ack into the meltin! pot1 We no% have three possiilities0 all eEually intri!uin!1
2 Pro$anochelys and Palaeochersis mi!ht e survivors of the land:d%ellin! animals that
had earlier sent some representatives to sea0 includin! the ancestors of !dontochelys1 This
hypothesis %ould su!!est that the shell evolved on land early0 and !dontochelys lost the carapace
in the %ater0 retainin! the ventral plastron1
3 The shell mi!ht have evolved in %ater0 as the Chinese authors su!!est0 %ith the plastron
over the elly evolvin! first0 and the carapace over the ack evolvin! later1 #n this case0 %hat do %e
make of Pro$anochelys and Palaeochersis0 %ho lived on land after !dontochelys lived0 %ith its
half shell0 in %aterB Pro$anochelys and Palaeochersis mi!ht have evolved the shell independently1
But there is another possiilityA
4 Pro$anochelys and Palaeochersis mi!ht represent an earlier return from the %ater to the
land1 #sn(t that a startlin!ly excitin! thou!htB
We are already pretty confident of the remarkale fact that the turtles accomplished an
evolutionary doulin! ack to the landA an early marEue of land (tortoises( %ent ack to the %atery
environment of their even earlier fish ancestors0 ecame sea turtles0 then returned to the land yet
a!ain0 as a ne% incarnation of land tortoises0 the Testudinidae1 That %e kno%0 or are nearly certain
of1 But no% %e are facin! up to the additional su!!estion that this doulin! ack happened twice>
*ot Hust to spa%n the modern tortoises0 ut much lon!er a!o0 to !ive rise to Pro$anochelys and
Palaeochersis in the Triassic1
#n another ook # descried '*& as (the Genetic Book of the 'ead(1 Because of the %ay
natural selection %orks0 there is a sense in %hich the '*& of an animal is a textual description of
the %orlds in %hich its ancestors %ere naturally selected1 ,or a fish0 the !enetic ook of the dead
descries ancestral seas1 ,or us and most mammals0 the early chapters of the ook are all set in the
sea and the later ones all out on land1 ,or %hales0 du!on!s0 marine i!uanas0 pen!uins0 seals0 sea
lions and turtles0 there is a third section of the ook %hich recounts their epic return to the provin!
!rounds of their remote past0 the sea1 But for the land tortoises0 perhaps t%ice independently on t%o
%idely separated occasions0 there is yet a fourth section of the ook devoted to a final : or is itB : re:
emer!ence0 yet a!ain to the land1 Can there e another animal for %hich the !enetic ook of the
dead is such a palimpsest of multiple evolutionary $:turnsB &s a partin! shot0 # cannot help
%onderin! aout those fresh%ater and rackish %ater forms 7(terrapins(80 %hich are close cousins of
the land tortoises1 'id their ancestors move directly from the sea into rackish and then fresh %aterB
'o they represent an intermediate sta!e on the %ay from the sea ack to the landB Or is it possile
that they constitute yet another doulin!:ack to the %ater from ancestors that %ere modern land
tortoisesB )ave the chelonians een shuttlin! ack and forth in evolutionary time et%een %ater
and landB Could the palimpsest e even more densely over:%ritten than # have so far su!!estedB
On 29 "ay 35590 as # %as correctin! the proofs of this ook0 a (missin! link( et%een lemur:
like and monkey:like primates %as announced in the online scientific Hournal P(!S !ne1 *amed
Darwini"s masillae0 it lived >@ million years a!o in rain forest in %hat is no% Germany1 #t is
claimed y the authors to e the most complete fossil primate ever foundA not Hust ones ut skin0
hair0 some internal or!ans and its last meal1 Beautiful as Darwini"s masillae undoutedly is 7see
colour pa!e 980 it comes trailin! clouds of hype that oscure clear thinkin!1 &ccordin! to Sky .ews
it is (the ei!hth %onder of the %orld( %hich (finally confirms Charles 'ar%in(s theory of evolution(1
Goodness meK The more:or:less nonsensical mystiEue of the (missin! link( seems to have lost none
of its po%er1
L "aHority opinion suspects the amateur palaeontolo!ist Charles 'a%son0 ut Stephen .ay
Gould intri!uin!ly floated the alternative theory that it mi!ht have een Pierre Teilhard de Chardin1
6ou may reco!niFe Teilhard(s name as the .esuit theolo!ian %hose later ook0 The Phenomenon of
%an0 %as to receive the !reatest ne!ative ook revie% of all time0 from the matchless Peter
"eda%ar 7reprinted in The Art of the Sol"ble and Pl"to's Rep"blic81
L #(m usin! (infinite( in the common0 often aused0 rhetorical sense of very very lar!e1 The
actual numer is the numer of pair%ise cominations of every species %ith every other : and that(s
as near infinite as makes no practical differenceK
L (Well:educated( reminds me of Peter "eda%ar(s %ickedly astute oservation that (the
spread of secondary and latterly of tertiary education has created a lar!e population of people0 often
%ith %ell:developed literary and scholarly tastes0 %ho have een educated far eyond their capacity
to undertake analytical thou!ht(1 #sn(t that pricelessB #t is the kind of %ritin! that makes me %ant to
rush out into the street to share %ith someody : anyody : ecause it is too !ood to keep to oneself1
L ,rom the term (clade(0 meanin! a !roup of or!anisms elieved to comprise all the
evolutionary descendants of a common ancestor1
O &t least accordin! to a consensus of Foolo!ists0 and # shall continue to use the irds0 for
the sake of ar!ument0 as an example of a !ood class1 Recent fossil research is sho%in! up a numer
of feathered dinosaurs0 and it is open to someody to claim that some of the modern animals %e call
irds are descended from a different !roup of feathered dinosaurs than others1 #f the most recent
common ancestor of all modern irds turns out to e an animal that %ould not e classified as a
ird0 # %ould have to revise my statement that the irds constitute a !ood class1
L #t(s een su!!ested0 y the %ay0 that this !i!antism %as made possile y the hi!her
oxy!en content in the atmosphere at that time1 #nsects lack lun!s0 and they reathe y means of tiny
air tues that pipe air throu!hout the ody1 &ir tues can(t mount such an intricately comprehensive
distriution system as lood tues can0 and it is plausile that this limits ody siFe1 That limit %ould
have een hi!her in an atmosphere %ith 4=S oxy!en0 instead of the mere 32S that %e reathe
today1 This provides a satisfyin! explanation for the !iant dra!onflies0 ut it may not necessarily e
the ri!ht one1 #ncidentally0 #(m puFFled %hy0 %ith so much oxy!en aout0 thin!s didn(t urst into
flames all the time1 Perhaps they did1 ,orest fires must have een more common than today0 and the
fossils indicate a hi!h incidence of fire:resistant plant species1 #t is not certain %hy the oxy!en
content of the atmosphere peaked durin! the Caroniferous and Permian1 #t may e associated %ith
the seEuesterin! of so much caron under the !round0 as coal1
L &n Oxford don of the old school0 %ho elieved he %as there to teach under!raduates0 he
%ould not have survived in today(s research:assessment culture1 With scarcely a sin!le pulished
article to his name0 his le!acy rests in the !enerations of !rateful pupils to %hom he imparted his
%isdom and at least some of his immense learnin!1
L This seems to e correct1 The !xford Dictionary of ?"otations su!!ests that the
commonly Euoted (seas( stems from a misprint in "asefield(s ori!inal 2953 editionA a nice example
of a successful mutant meme1
L #(m advised that this doesn(t make a lot of sense in Greek1 #f it %ere Pro$onochelys it
%ould make perfect sense1 #t %ould mean somethin! like (ancestral tortoise( or (primeval tortoise(0
and # can(t help feelin! that that may e %hat the ori!inal authors intended %hen they named it1
$nfortunately0 the rules of Foolo!ical nomenclature are strict0 and even ovious mistakes can(t e
chan!ed0 once they are enshrined in a namin! pulication1 The taxonomy is littered %ith such
fossiliFed mistakes1 "y favourite is /haya0 &frican maho!any1 -e!end 7%hich # lon! to elieve8
has it that in a local lan!ua!e it means (# don(t kno%(0 %ith the presumed sutext0 (&nd # don(t care
and %hy don(t you stop askin! stupid Euestions aout plant names1(
' &RW#*(S treatment of human evolution in his most famous %ork0 !n the !ri$in of
Species0 is limited to t%elve portentous %ordsA (-i!ht %ill e thro%n on the ori!in of man and his
history1( That is the %ordin! in the first edition0 %hich is the edition # al%ays cite unless other%ise
stated1 By the sixth 7and last8 edition0 'ar%in allo%ed himself to stretch a point0 and the sentence
ecame ("uch li!ht %ill e thro%n on the ori!in of man and his history1( # like to think of his pen0
poised over the fifth edition0 %hile the !reat man Hudiciously pondered %hether to indul!e himself
in the luxury of ("uch(1 Even %ith it0 the sentence is a calculated understatement1
'ar%in delierately deferred his treatment of human evolution to another ook0 The
Descent of %an1 Perhaps it is not surprisin! that the t%o volumes of that later %ork devote more
space to the topic of its sutitle0 Selection in Relation to Sex 7investi!ated lar!ely in irds80 than to
human evolution1 *ot surprisin! ecause0 at the time of 'ar%in(s %ritin!0 there %ere no fossils at all
linkin! us to our closest relatives amon! the apes1 'ar%in had only livin! apes to look at0 and he
used them %ell0 ar!uin! correctly 7and almost alone8 that our closest livin! relatives %ere all
&frican 7!orillas and chimpanFees : onoos %ere not reco!niFed as separate from chimpanFees in
those days0 ut they are &frican too80 and therefore predictin! that0 if ancestral human fossils %ere
ever to e found0 &frica %as the place to search1 'ar%in re!retted the paucity of fossils0 ut he
maintained a roustly ullish attitude to it1 Citin! -yell0 his mentor and the !reat !eolo!ist of the
time0 he pointed out that (in all the verterate classes the discovery of fossil remains has een an
extremely slo% and fortuitous process( and added0 (*or should it e for!otten that those re!ions
%hich are the most likely to afford remains connectin! man %ith some extinct ape:like creature0
have not as yet een searched y !eolo!ists1( )e meant &frica0 and the Euest %as not helped y the
fact that his immediate successors lar!ely i!nored his advice and searched &sia instead1
#t %as indeed in &sia that the (missin! links( first e!an to ecome less missin!1 But those
first fossils to e discovered %ere relatively recent0 less than a million years old0 datin! from a time
%hen hominids %ere pretty close to modern humans and had mi!rated out of &frica and reached the
,ar East1 They %ere called (.ava "an( and (Pekin! "an( after their discovery sites1L .ava "an %as
discovered y the 'utch anthropolo!ist Eu!ene 'uois in 2;921 )e named it Pithecanthrop"s
erect"s0 si!nifyin! his elief that he had realiFed his life(s amition and found (the missin! link(1
'isa!reement came from t%o opposite sources0 %hich rather proved his pointA some said his fossil
%as purely human0 others that it %as a !iant !ion1 -ater in his rather emittered and cantankerous
life0 'uois resented the su!!estion that the more recently discovered Pekin! fossils %ere similar to
his .ava "an1 ,iercely possessive aout0 not to say protective of0 his fossil0 'uois elieved that
only .ava "an %as the true missin! link1 To emphasiFe the distinction from the various Pekin! "an
fossils0 he descried them as far closer to modern man0 and his o%n .ava "an of Trinil as
intermediate et%een man and ape1
Pithecanthrop"s M.ava "anN %as not a man0 ut a !i!antic !enus allied to the !ions0
ho%ever superior to the !ions on account of its exceedin!ly lar!e rain volume and distin!uished
at the same time y its faculty of assumin! an erect attitude and !ait1 #t had the doule cephaliFation
Mratio of rain siFe to ody siFeN of the anthropoid apes in !eneral and half that of man 1 1 1
#t %as the surprisin! volume of the rain : %hich is very much too lar!e for an anthropoid
ape0 and %hich is small compared %ith the avera!e0 thou!h not smaller than the smallest human
rain : that led to the no% almost !eneral vie% that the (&pe "an( of Trinil0 .ava %as really a
primitive "an1 "orpholo!ically0 ho%ever0 the calvaria MskullcapN closely resemles that of
anthropoid apes0 especially the !ion 1 1 1
#t can(t have improved 'uois( temper that others took him to e sayin! that
Pithecanthrop"s %as Hust a !iant !ion0 not intermediate et%een them and humans at all0 and he
%as at pains to reassert his earlier standA (# still elieve0 no% more firmly than ever0 that the
Pithecanthrop"s of Trinil is the real Qmissin! linkQ1(
Creationists have from time to time used as a political %eapon the alle!ation that 'uois
acked off from his claim that Pithecanthrop"s %as an intermediate ape:man1 The creationist
or!aniFation &ns%ers in Genesis has0 ho%ever0 added it to their list of discredited ar!uments %hich
they no% say should not e used1 #t is to their credit that they maintain such a list at all1 &s # said0
oth the .ava and Pekin! specimens of Pithecanthrop"s have no% een sho%n to e Euite youn!0
less than a million years old1 They are no% classified alon! %ith us in the !enus )omo0 retainin!
'uois( specific name erect"sA )omo erect"s1
'uois chose the %ron! part of the %orld for his sin!le:minded Euest for the (missin! link(1
#t %as natural for a 'utchman to head first for the 'utch East #ndies0 ut a man of his dedication
should have follo%ed 'ar%in(s advice and !one on to &fricaA for &frica is %here our ancestors
evolved0 as %e shall see1 So %hat %ere these )omo erect"s specimens doin! out of &fricaB The
phrase (out of &frica( has een orro%ed from Daren Blixen L to refer to the !reat exodus of our
ancestors from &frica1 But there %ere t%o exoduses and it is important not to confuse them1
Relatively recently0 maye less than 2550555 years a!o0 rovin! ands of )omo sapiens lookin!
pretty much like us left &frica and diversified into all the races that %e see around the %orld todayA
#nuit0 native &mericans0 native &ustralians0 Chinese0 and so on1 #t is to this recent exodus that the
phrase (out of &frica( is normally applied1 But there %as an earlier exodus from &frica0 and these
erect"s pioneers left fossils in &sia and Europe0 includin! the .ava and Pekin! specimens1 The
oldest fossil kno%n outside &frica %as found in the central &sian country of Geor!ia and dued
(Geor!ian "an(A a diminutive creature %hose 7rather %ell:preserved8 skull is dated0 y modern
methods0 to aout 21; million years a!o1 #t has een called )omo $eor$ic"s 7y some taxonomists0
althou!h others don(t reco!niFe it as a separate species8 to indicate that it seems rather more
primitive than the rest of the early refu!ees from &frica0 %ho are all classified as )omo erect"s1
Some stone tools sli!htly older than Geor!ian "an have Hust een discovered in "alaysia0 sparkin!
a ne% search for fossil ones in that peninsula1 But in any case0 all these early &sian fossils are
pretty close to modern humans and all are no%adays classified in the !enus )omoJ for our earlier
antecedents %e must !o to &frica1 ,irst0 thou!h0 let(s pause to ask %hat %e should expect of a
(missin! link(1
Homo georgicus
Suppose0 for the sake of ar!ument0 %e take seriously the ori!inal confused meanin! of the
term (missin! link(0 and seek an intermediate et%een chimpanFees 7see ri!ht8 and ourselves1 We
are not descended from chimpanFees0 ut it is a fair et that the common ancestor that %e share
%ith them %as more like a chimp than like us1 #n particular0 it didn(t have a hu!e rain like ours0 it
proaly didn(t %alk upri!ht as %e do0 it proaly %as a lot hairier than %e are0 and it surely didn(t
have such advanced human features as lan!ua!e1 So0 even thou!h %e must remain adamant0 in the
face of common misunderstandin!0 that %e are not descended from chimpanFees0 there(s still no
harm in askin! %hat an intermediate et%een somethin! like a chimpanFee and us %ould look like1
Well0 hair and lan!ua!e don(t fossiliFe %ell0 ut %e can !et !ood clues aout rain siFe from
the skull0 and !ood clues aout !ait from the %hole skeleton 7includin! the skull0 for the foramen
ma$n"m0 the hole for the spinal cord0 points do%n%ards in ipeds0 more ack%ards in Euadrupeds81
Possile candidates for missin! links mi!ht have any of the follo%in! attriutesA
2 #ntermediate rain siFe and intermediate !aitA perhaps a sort of stoopin! shamle rather
than the proudly erect earin! favoured y ser!eant maHors and deportment mistresses1
3 ChimpanFee:siFed rain %ith human upri!ht !ait1
4 -ar!e0 more human:like rain0 %alkin! on all fours like a chimp1
So0 earin! these possiilities in mind0 let(s examine some of the many &frican fossils that
are no% availale to us0 ut unfortunately %ere not availale to 'ar%in1
#(" ST#-- "#SC)#E+O$S-6 )OP#*G 1 1 1
"olecular evidence 7%hich # shall come on to in Chapter 258 sho%s that the common
ancestor %e share %ith chimpanFees lived aout six million years a!o or a it earlier0 so let(s split
the difference and look at some three:million:year:old fossils1 The most famous fossil of this
vinta!e is (-ucy(0 classified y her discoverer in Ethiopia0 'onald .ohanson0 as A"stralopithec"s
afarensis1 $nfortunately %e have only fra!ments of -ucy(s cranium0 ut her lo%er Ha% is unusually
%ell preserved1 She %as small y modern standards0 althou!h not as small as )omo floresiensis0 the
tiny creature the ne%spapers have irritatin!ly dued (the )oit(0 %hich died out tantaliFin!ly
recently on the island of ,lores in #ndonesia1 -ucy(s skeleton is complete enou!h to su!!est that she
%alked upri!ht on the !round0 ut proaly also sou!ht refu!e in trees0 %here she %as an a!ile
climer1 There is !ood evidence that the ones attriuted to -ucy really did all come from a sin!le
individual1 The same is not true of the so:called (,irst ,amily(0 a collection of ones from at least
thirteen individuals0 similar to -ucy and of approximately the same vinta!e0 %ho someho% ecame
uried to!ether0 also in Ethiopia1 The fra!ments from -ucy and from the ,irst ,amily !ive a !ood
idea of %hat A"stralopithec"s afarensis looked like0 ut it is hard to make an authentically complete
reconstruction from pieces of many different individuals1 ,ortunately0 a rather complete skull
kno%n as &- >>>:3 7ri!ht8 %as discovered in 2993 in the same area of Ethiopia0 and this confirmed
the tentative reconstructions that had previously een made1
The conclusion from studies of -ucy and her kind is that they had rains aout the same siFe
as chimpanFees( ut0 unlike chimpanFees0 they %alked upri!ht on their hind le!s0 as %e do : the
second of our three hypothetical scenarios1 (-ucys( %ere a it like upri!ht:%alkin! chimps1 Their
ipedality is dramatically confirmed y the poi!nantly evocative set of footprints discovered y
"ary -eakey in fossiliFed volcanic ash1 These are further south0 at -aetoli in TanFania0 and they are
older than -ucy and &- >>>:3A aout 41< million years1 They are usually attriuted to a pair of
A"stralopithec"s afarensis %alkin! to!ether 7hand in handB8 ut %hat matters is that0 y 41< million
years a!o0 an erect ape %alked the Earth0 on t%o feet %hich %ere pretty much like ours althou!h its
rain %as the siFe of a chimpanFee(s1
A1 222)2
#t seems Euite likely that the species %e call A"stralopithec"safarensis: -ucy(s species :
included our ancestors of three million years a!o1 Other fossils have een placed in different species
of the same !enus0 and it is virtually certain that our ancestors %ere memers of that !enus1 The
first &ustralopithecine to e discovered0 and the type specimen of the !enus0 %as the so:called
Taun! Child1 &t the a!e of three and a half the Taun! Child %as eaten y an ea!le1 The evidence is
that dama!e marks to the eye sockets of the fossil are identical to marks made y modern ea!les on
modern monkeys as they rip out their eyes1 Poor little Taun! Child0 shriekin! on the %ind as you
%ere orne aloft y the aEuiline fury0 you %ould have found no comfort in your destined fame0 t%o
and a half million years on0 as the type specimen of A"stralopithec"s african"s1 Poor Taun!
mother0 %eepin! in the Pliocene1
The type specimen is the first individual of a ne% species to e named and officially !iven
the vir!in lael in a museum1 Theoretically0 later finds are compared a!ainst the type specimen to
see if they match1 The Taun! Child %as discovered and !iven rand ne% !enus and species names
y the South &frican anthropolo!ist Raymond 'art in 293>1
What(s the difference et%een (species( and (!enus(B -et(s !et the Euestion s%iftly out of the
%ay0 efore proceedin!1 Genus is the more inclusive division1 & species elon!s %ithin a !enus0
and often it shares the !enus %ith other species1 )omo sapiens and )omo erect"s are t%o species
%ithin the !enus )omo4 A"stralopithec"s african"s and A"stralopithec"s afarensis are t%o species
%ithin the !enus A"stralopithec"s1 The -atin name of an animal or plant al%ays includes a !eneric
name 7%ith an initial capital letter8 follo%ed y a specific name 7%ithout a capital letter81 Both
names are %ritten in italics1 Sometimes there is an additional su:specific name0 %hich follo%s the
specific name0 as in0 for example0 )omo sapiens neanderthalensis1 Taxonomists often dispute
names1 "any0 for example0 %ould speak of )omo neanderthalensis not )omo sapiens
neanderthalensis0 elevatin! *eanderthal man from su:species to species status1 Generic names and
specific names are also often disputed0 and often chan!e %ith successive revisions in the scientific
literature1 Paranthrop"s boisei has een0 in its time0 @in6anthrop"s boisei and A"stralopithec"s
boisei0 L and is still often referred to0 informally0 as a roust &ustralopithecine : as opposed to the
t%o (!racile( 7slender8 species of A"stralopithec"s mentioned aove1 One of the main messa!es of
this chapter concerns the some%hat aritrary nature of Foolo!ical classification1
Raymond 'art0 then0 !ave the name A"stralopithec"s to the Taun! Child0 the type specimen
of the !enus0 and %e have een stuck %ith this depressin!ly unima!inative name for our ancestor
ever since1 #t simply means (southern ape(1 *othin! to do %ith &ustralia0 %hich Hust means (southern
country(1 6ou(d think 'art mi!ht have thou!ht of a more ima!inative name for such an important
!enus1 )e mi!ht even have !uessed that other memers of the !enus %ould later e discovered
north of the eEuator1
Sli!htly older than the Taun! Child0 one of the most eautifully preserved skulls %e have0
althou!h lackin! a lo%er Ha%0 is called ("rs Ples(1 "rs Ples0 %ho may actually have een a small
male rather than a lar!e female0 otained (her( nickname ecause she %as ori!inally classified in the
!enus Plesianthrop"s1 This means (nearly human(0 %hich is a etter name than (southern ape(1 One
mi!ht have hoped that0 %hen later taxonomists decided that "rs Ples and her kind %ere really of the
same !enus as the Taun! Child0 Plesianthrop"s %ould have ecome the name for all of them1
$nfortunately0 the rules of Foolo!ical nomenclature are strict to the point of pedantry1 Priority of
namin! takes precedence over sense and suitaility1 (Southern ape( mi!ht e a lousy name ut no
matterA it predates the much more sensile Plesianthrop"s and %e seem to e stuck %ith it0 unless 1 1
1 #(m still mischievously hopin! someody %ill uncover0 in a dusty dra%er in a South &frican
museum0 a lon!:for!otten fossil0 clearly the same kind as "rs Ples and the Taun! Child0 ut earin!
the scra%led lael0 ( )emianthrop"s type specimen0 2935(1 &t a stroke0 all the museums in the %orld
%ould immediately have to relael their A"stralopithec"s specimens and casts0 and all ooks and
articles on hominid prehistory %ould have to follo% suit1 Wordprocessin! pro!rams across the
%orld %ould %ork overtime sniffin! out any occurrences of A"stralopithec"s and replacin! them
%ith )emianthrop"s1 # can(t think of any other case %here international rules are potent enou!h to
dictate a %orld%ide and ackdated chan!e of lan!ua!e overni!ht1
'4rs 7les'
*o% for my next important point aout alle!edly missin! links and the aritrariness of
names1 Oviously0 %hen "rs Ples(s name %as chan!ed from Plesianthrop"s to A"stralopithec"s0
nothin! chan!ed in the real %orld at all1 Presumaly noody %ould e tempted to think anythin!
else1 But consider a similar case %here a fossil is re:examined and moved0 for anatomical reasons0
from one !enus to another1 Or %here its !eneric status is disputed : and this very freEuently happens
: y rival anthropolo!ists1 #t is0 after all0 essential to the lo!ic of evolution that there must have
existed individuals sittin! exactly on the orderline et%een t%o !enera0 say A"stralopithec"s and
)omo1 #t is easy to look at "rs Ples and a modern )omo sapiens skull and say0 yes0 there is no
dout these t%o skulls elon! in different !enera1 #f %e assume0 as almost every anthropolo!ist
today accepts0 that all memers of the !enus )omo are descended from ancestors elon!in! to the
!enus %e call A"stralopithec"s0 it necessarily follo%s that0 some%here alon! the chain of descent
from one species to the other0 there must have een at least one individual %ho sat exactly on the
orderline1 This is an important point0 so let me stay %ith it a little lon!er1
KN4 5R 191:
KN4 5R 12;<
Bearin! in mind the shape of "rs Ples(s skull as a representative of A"stralopithec"s
african"s 31< million years a!o0 have a look at the top skull opposite0 called D*" ER 2;241 Then
look at the one underneath it0 called D*" ER 2>@51 Both are dated at approximately 219 million
years a!o0 and oth are placed y most authorities in the !enus )omo1 Today0 2;24 is classified as
)omo habilis0 ut it %asn(t al%ays1 $ntil recently0 2>@5 %as too0 ut there is no% a move afoot to
reclassify it as )omo r"dolfensis1 Once a!ain0 see ho% fickle and transitory our names are1 But no
matterA oth have an apparently a!reed foothold in the !enus )omo1 The ovious difference from
"rs Ples and her kind is that she had a more for%ard:protrudin! face and a smaller rain:case1 #n
oth respects0 2;24 and 2>@5 seem more human0 "rs Ples more (ape:like(1
*o% look at the skull elo%0 called (T%i!!y(1 T%i!!y is also normally classified no%adays
as )omo habilis1 But her for%ard:pointin! muFFle has more of a su!!estion of "rs Ples aout it
than of 2>@5 or 2;241 6ou %ill perhaps not e surprised to e told that T%i!!y has een placed y
some anthropolo!ists in the !enus A"stralopithec"s and y other anthropolo!ists in )omo1 #n fact0
each of these three fossils has een0 at various times0 classified as )omo habilis and as
A"stralopithec"s habilis1 &s # have already noted0 some authorities at some times have !iven 2>@5 a
different specific name0 chan!in! habilis to r"dolfensis1 &nd0 to cap it all0 the specific name
r"dolfensis has een fastened to oth !eneric names0 A"stralopithec"s and )omo1 #n summary0
these three fossils have een variously called0 y different authorities at different times0 the
follo%in! ran!e of namesA
D*" ER 2;24A
A"stralopithec"s habilis0 )omo habilis
D*" ER 2>@5A
A"stralopithec"s habilis0 )omo habilis0
A"stralopithec"s r"dolfensis0 )omo r"dolfensis
O) 3> 7(T%i!!y(8A
A"stralopithec"s habilis0 )omo habilis
Should such a confusion of names shake our confidence in evolutionary scienceB Puite the
contrary1 #t is exactly %hat %e should expect0 !iven that these creatures are all evolutionary
intermediates0 links that %ere formerly missin! ut are missin! no lon!er1 We should e positively
%orried if there %ere no intermediates so close to orderlines as to e difficult to classify1 #ndeed0
on the evolutionary vie%0 the conferrin! of discrete names should actually ecome impossile if
only the fossil record %ere more complete1 #n one %ay0 it is fortunate that fossils are so rare1 #f %e
had a continuous and unroken fossil record0 the !rantin! of distinct names to species and !enera
%ould ecome impossile0 or at least very prolematical1 #t is a fair conclusion that the predominant
source of discord amon! palaeoanthropolo!ists : %hether such and such a fossil elon!s in this
speciesG!enus or that : is deeply and interestin!ly futile1
)old in your head the hypothetical notion that %e mi!ht0 y some fluke0 have een lessed
%ith a continuous fossil record of all evolutionary chan!e0 %ith no links missin! at all1 *o% look at
the four -atin names that have een applied to 2>@51 On the face of it0 the chan!e from habilis to
r"dolfensis %ould seem to e a smaller chan!e than the one from A"stralopithec"s to )omo1 T%o
species %ithin a !enus are more like each other than t%o !enera1 &ren(t theyB #sn(t that the %hole
asis for the distinction et%een the !enus level 7say )omo or Pan as alternative !enera of &frican
apes8 and the species level 7say tro$lodytes or panisc"s %ithin the chimpanFees8 in the hierarchy of
classificationB Well0 yes0 that is ri!ht %hen %e are classifyin! modern animals0 %hich can e
thou!ht of as the tips of the t%i!s on the evolutionary tree0 %ith their antecedents on the inside of
the tree(s cro%n all comfortaly dead and out of the %ay1 *aturally0 those t%i!s that Hoin each other
further ack 7further into the interior of the tree(s cro%n8 %ill tend to e less alike than those %hose
Hunction 7more recent common ancestor8 is nearer the tips1 The system %orks0 as lon! as %e don(t
try to classify the dead antecedents1 But as soon as %e include our hypothetically complete fossil
record0 all the neat separations reak do%n1 'iscrete names ecome0 as a !eneral rule0 impossile to
apply1 We can easily see this if %e %alk steadily ack%ards throu!h time0 much as %e did %ith the
raits in Chapter 31
&s %e trace the ancestry of modern )omo sapiens ack%ards0 there must come a time %hen
the difference from livin! people is sufficiently !reat to deserve a different specific name0 say
)omo er$aster1 6et0 every step of the %ay0 individuals %ere presumaly sufficiently similar to their
parents and their children to e placed in the same species1 *o% %e !o ack further0 tracin! the
ancestry of )omo er$aster0 and there must come a time %hen %e reach individuals %ho are
sufficiently different from (mainstream( er$aster to deserve a different specific name0 say )omo
habilis1 &nd no% %e come to the point of this ar!ument1 &s %e !o ack further still0 at some point
%e must start to hit individuals sufficiently different from modern )omo sapiens to deserve a
different !enus nameA say A"stralopithec"s1 The troule is0 (sufficiently different from modern
)omo sapiens( is another matter entirely from (sufficiently different from the earliest )omo(0 here
desi!nated )omo habilis1 Think aout the first specimen of )omo habilis to e orn1 )er parents
%ere A"stralopithec"s1 She elon!ed to a different !enus from her parentsB That(s Hust dopeyK 6es
it certainly is1 But it is not reality that(s at fault0 it(s our human insistence on shovin! everythin! into
a named cate!ory1 #n reality0 there %as no such creature as the first specimen of )omo habilis1
There %as no first specimen of any species or any !enus or any order or any class or any phylum1
Every creature that has ever een orn %ould have een classified : had there een a Foolo!ist
around to do the classifyin! : as elon!in! to exactly the same species as its parents and its
children1 6et0 %ith the hindsi!ht of modernity0 and %ith the enefit : yes0 in this one paradoxical
sense benefit : of the fact that most of the links are missin!0 classification into distinct species0
!enera0 families0 orders0 classes and phyla ecomes possile1
# %ish %e really did have a complete and unroken trail of fossils0 a cinematic record of all
evolutionary chan!e as it happened1 # %ish it0 not least ecause #(d love to see the e!! all over the
faces of those Foolo!ists and anthropolo!ists %ho en!a!e in lifelon! feuds %ith each other over
%hether such and such a fossil elon!s to this species or that0 this !enus or that1 Gentlemen : #
%onder %hy it never seems to e ladies : you are ar!uin! aout %ords0 not reality1 &s 'ar%in
himself said0 in The Descent of %an0 (#n a series of forms !raduatin! insensily from some apelike
creature to man as he no% exists0 it %ould e impossile to fix on any definite point %here the term
QmanQ ou!ht to e used1(
-et(s move on throu!h the fossils0 and look at some more recent links amon! those that are
no lon!er missin!0 althou!h they %ere missin! in 'ar%in(s time1 What intermediates can %e find
et%een ourselves and the various creatures like 2>@5 and T%i!!y0 %ho are sometimes called
)omo and sometimes called A"stralopithec"sB We(ve already met some of them0 as .ava "an and
Pekin! "an0 normally classified as )omo erect"s1 But those t%o lived in &sia0 and there(s !ood
evidence that most of our human evolution took place in &frica1 .ava "an and Pekin! "an and
their kind %ere emi!rants from the mother continent of &frica1 Within &frica itself0 their
eEuivalents are no%adays usually classified as )omo er$aster0 althou!h for many years they %ere
all called )omo erect"s : yet another illustration of the fickleness of our namin! procedures1 The
most famous specimen of )omo er$aster0 and one of the most complete pre:human fossils ever
found0 is the Turkana Boy0 or *ariokotome Boy0 discovered y Damoya Dimeu0 star fossil:finder of
Richard -eakey(s team of palaeontolo!ists1
Homo erectus
The Turkana Boy lived approximately 21< million years a!o and died at the a!e of aout
eleven1 There are indications that he %ould have !ro%n to a hei!ht of < feet if he had lived to
adulthood1 )is proHected adult rain volume %ould have een aout 955 cuic centimetres 7cc81
This %as typical of )omo er$asterG erect"s rains0 %hich varied around 20555 cc1 #t is si!nificantly
smaller than modern human rains0 %hich vary around 20455 or 20>55 cc0 ut lar!er than )omo
habilis 7around <55 cc8 %hich in turn %as lar!er than A"stralopithec"s 7around >55 cc8 and
chimpanFees 7around the same81 6ou(ll rememer %e concluded that our ancestor of three million
years a!o had the rain of a chimpanFee ut %alked on its hind le!s1 ,rom this %e mi!ht presume
that the second half of the story0 from 4 million years a!o to recent times0 %ould e a tale of
increasin! rain siFe1 &nd so0 indeed0 it proves1
)omo er$asterAerect"s0 of %hich %e have many fossil specimens0 is a very persuasive
half%ay link0 no lon!er missin!0 et%een )omo sapiens today and )omo habilis t%o million years
a!o0 %hich is in turn a eautiful link ack to A"stralopithec"s three million years a!o0 %hich0 as %e
sa%0 could pretty %ell e descried as an upri!ht:%alkin! chimpanFee1 )o% many links do you
need0 efore you concede that they are no lon!er (missin!(B &nd can %e also rid!e the !ap et%een
)omo er$aster and modern )omo sapiensB 6esA %e have a rich lode of fossils0 coverin! the last
fe% hundred thousand years0 %hich are intermediate et%een them1 Some have een !iven species
names0 like )omo heidelber$ensis0 )omo rhodesiensis and )omo neanderthalensis1 Others 7and
sometimes the same ones8 are called (archaic( )omo sapiens1 But0 as # keep repeatin!0 names don(t
matter1 What matters is that the links are no lon!er missin!1 #ntermediates aound1
.$ST GO &*' -OOD
So0 %e have fine fossil documentation of !radual chan!e0 all the %ay from -ucy0 the
(upri!ht:%alkin! chimp( of three million years a!o0 to ourselves today1 )o% do history:deniers cope
%ith this evidenceB Some y literal denial1 # encountered this in an intervie% # did for the Channel
,our television documentary The Geni"s of #harles Darwin in 355;1 # %as intervie%in! Wendy
Wri!ht0 President of (Concerned Women for &merica(1 )er opinion that (The mornin!:after pill is a
pedophile(s est friend( !ives a fair idea of her po%ers of reasonin!0 and she fully lived up to
expectation durin! our intervie%1 Only a very small part of the intervie% %as used for the television
documentary1 What follo%s is a much fuller transcript0 ut oviously for the purposes of this
chapter # have confined myself to those places %here %e discussed the fossil record of human
WendyA What # !o ack to is the evolutionists are still lackin! the science to ack it up1 But
instead %hat happens is science that doesn(t olster the case for evolution !ets censored out1 Such as
there is no evidence of evolution from !oin! from one species to another species1 #f that0 if
evolution had occurred then surely %hether it(s !oin! from irds to mammals or0 or0 even eyond
that surely there(d e at least one evidence1
RichardA There(s a massive amount of evidence1 #(m sorry ut you people keep repeatin! that
like a kind of mantra ecause you0 you0 Hust listen to each other1 # mean0 if only you %ould Hust open
your eyes and look at the evidence1
WendyA Sho% it to me0 sho% me the0 sho% me the ones0 sho% me the carcass0 sho% me the
evidence of the in:et%een sta!es from one species to another1
RichardA Every time a fossil is found %hich is in et%een one species and another you !uys
say0 (&h0 no% %e(ve !ot t%o !aps %here there0 %here previously there %as only one1( # mean almost
every fossil you find is intermediate et%een somethin! and somethin! else1
Wendy M la"$hsNA #f that %ere the case0 the Smithsonian *atural )istory "useum %ould e
filled %ith these examples ut it isn(t1
RichardA #t is0 it is 1 1 1 in the case of humans0 since 'ar%in(s time there(s no% an enormous
amount of evidence aout intermediates in human fossils and you(ve !ot various species of
A"stralopithec"s for example0 and 1 1 1 then you(ve !ot )omo habilis : these are intermediates
et%een A"stralopithec"s %hich %as an older species and )omo sapiens %hich is a youn!er
species1 # mean0 %hy don(t you see those as intermediatesB
WendyA 1 1 1 if evolution has had the actual evidence then it %ould e displayed in museums
not Hust in illustrations1
RichardA # Hust told you aout A"stralopithec"s0 )omo habilis0 )omo erect"s0 )omo sapiens
: archaic )omo sapiens and then modern )omo sapiens : that(s a eautiful series of intermediates1
WendyA 6ou(re still lackin! the material evidence so 1 1 1
RichardA The material evidence is there1 Go to the museum and look at it 1 1 1 # don(t have
them here oviously0 ut you can !o to any museum and you can see A"stralopithec"s0 you can see
)omo habilis0 you can see )omo erect"s0 you can see archaic )omo sapiens and modern )omo
sapiens1 & eautiful series of intermediates1 Why do you keep sayin! (Present me %ith the evidence(
%hen #(ve done soB Go to the museum and look1
WendyA &nd # have1 # have !one to the museums and there are so many of us %ho still are
not convinced 1 1 1
RichardA )ave you seen0 have you seen )omo erect"sB
WendyA &nd # think there(s this effort0 this rather a!!ressive effort to try and talk over us and
to censor us1 #t seems to come out of a frustration that so many people still don(t elieve in
evolution1 *o% if evolutionists %ere so confident in their eliefs there %ouldn(t e the effort to
censor out information1 #t sho%s that evolution is still lackin! and is Euestionale1
RichardA # am 1 1 1 # confess to ein! frustrated1 #t(s not aout suppression0 it(s aout the fact
that # have told you aout four or five fossils 1 1 1 M Wendy la"$hsN 1 1 1 and you seem to simply e
i!norin! %hat #(m sayin! 1 1 1 Why don(t you !o and look at those fossilsB
WendyA 1 1 1 #f they %ere in the museums %hich #(ve een to many times0 then # %ould look at
them oHectively0 ut %hat # !o ack to is 1 1 1
RichardA They are in the museum1
WendyA What # !o ack to is that the philosophy of evolution can lead to ideolo!ies that
have een so destructive to the human race 1 1 1
RichardA 6es0 ut %ouldn(t it e a !ood idea0 instead of pointin! to misperceptions of
'ar%inism0 %hich have een hideously misused politically0 if you tried to understand 'ar%inism0
then you(d e in a position to counteract these horrile misunderstandin!s1
WendyA Well actually %e are so often forced y the a!!ressiveness of those %ho favour
evolution1 #t(s not as if %e are hidden from this information that you keep presentin!1 #t(s not as if it
is unkno%n to us0 ecause %e can(t !et a%ay from it1 #t(s pushed on us all the time1 But # think your
frustration comes from the fact that so many of us %ho have seen your information still don(t uy
into your ideolo!y1
RichardA )ave you seen )omo erect"sB )ave you seen )omo habilisB )ave you seen
A"stralopithec"sB #(ve asked you that Euestion1
WendyA What #(ve seen is that in the museums and in the textooks %henever they claim to
sho% the evolutionary differences from one species to another0 it relies on illustrations and
dra%in!s 1 1 1 not any material evidence1
RichardA Well0 you mi!ht have to !o to the *airoi "useum to see the ori!inal fossils ut
you can see casts of fossils : exact copies of these fossils in any maHor museum you care to look at1
WendyA Well0 let me ask you %hy you are so a!!ressiveB Why is it so important to you that
everyone elieves like you elieveB
RichardA #(m not talkin! aout elief0 #(m talkin! aout facts1 #(ve told you aout certain
fossils0 and every time # ask you aout them you evade the Euestion and turn to somethin! else1
WendyA 1 1 1 There should e over%helmin! tons of material evidence not Hust an isolated
thin!0 ut a!ain0 there is not evidence1
RichardA # happened to pick hominid fossils ecause # thou!ht you(d e most interested in
them0 ut you can find similar fossils from any verterate !roup you care to name1
WendyA But # !uess # !o ack to %hy is it so important to you that everyone elieves in
evolution 1 1 1
RichardA # don(t like the %ord elief1 # prefer to Hust ask people to look at the evidence0 and
#(m askin! you to look at the evidence 1 1 1 # %ant you to !o to museums and look at the facts and
don(t elieve %hat you have een told that there is no evidence1 .ust !o and look at the evidence1
Wendy M la"$hsNA &nd yes0 and %hat # %ould say 1 1 1
RichardA #t(s not funny1 # mean0 really !o0 !o1 #(ve told you aout hominid fossils0 and you
can !o and see the evolution of the horse0 you can !o and look at the evolution of the early
mammals0 you can !o and look at the evolution of fish0 you can !o and look at the transition from
fish to land:livin! amphiians and reptiles1 &ny of those thin!s you %ill find in any !ood museum1
.ust open your eyes and look at the facts1
WendyA &nd # %ould say open your eyes and see the communities that have een uilt y
those %ho elieve in a lovin! God %ho created each one of us 1 1 1
#t mi!ht seem0 in that exchan!e0 that # %as ein! needlessly ostinate in hammerin! home
the reEuest that she should !o to a museum and look0 ut # really meant it1 These people have een
coached to say0 (There are no fossils0 sho% me the evidence0 sho% me Hust one fossil 1 1 1( and they
say it so often that they come to elieve it1 So # tried the experiment of mentionin! three or four
fossils to this %oman and not lettin! her !et a%ay %ith simply i!norin! them1 The results are
depressin!0 and a !ood example of the commonest tactic used y history:deniers %hen confronted
%ith the evidence of history : namely0 Hust i!nore it and repeat the mantraA (Sho% me the fossils1
Where are the fossilsB There are no fossils1 .ust sho% me one intermediate fossil0 that(s all # ask 1 1 1(
Others efuddle themselves %ith names0 and the inevitale tendency names have to make
false divisions %here there are none1 Every fossil that mi!ht potentially e intermediate is al%ays
classified as either )omo or A"stralopithec"s1 *one is ever classified as intermediate1 Therefore
there are no intermediates1 But0 as # have explained aove0 this is an inevitale conseEuence of the
conventions of Foolo!ical nomenclature0 not a fact aout the real %orld1 The most perfect
intermediate you could possily ima!ine %ould still find itself shoehorned into either )omo or
A"stralopithec"s1 #n fact0 it %ould proaly e called )omo y half the palaeontolo!ists and
A"stralopithec"s y the other half1 &nd unfortunately0 instead of !ettin! to!ether to a!ree that
ami!uously intermediate fossils are exactly %hat %e should expect on the evolution theory0 the
palaeontolo!ists could proaly e relied upon to !ive an entirely false impression y seemin!
almost to come to lo%s over their terminolo!ical disa!reement1
#t(s a it like the le!al distinction et%een an adult and a minor1 ,or le!al purposes0 and for
decidin! %hether a youn! person is old enou!h to vote or Hoin the army0 it is necessary to make an
asolute distinction1 #n 29<9 the le!al votin! a!e in Britain %as lo%ered from t%enty:one to
ei!hteen 7in 29@2 the same chan!e %as made in the $S&81 *o% there(s talk of lo%erin! it to
sixteen1 But0 %hatever the le!al votin! a!e may e0 noody seriously thinks the stroke of midni!ht
on the ei!hteenth 7or t%enty:first0 or sixteenth8 irthday actually turns you into a different kind of
person1 *oody seriously elieves there are t%o kinds of people0 children and adults0 %ith (no
intermediates(1 Oviously %e all understand that the %hole period of !ro%in! up is one lon!
exercise in intermediacy1 Some of us0 it mi!ht e said0 have never really !ro%n up1 Similarly0
human evolution0 from somethin! like A"stralopithec"s afarensis to )omo sapiens0 consisted of an
unroken series of parents !ivin! irth to children %ho %ould certainly have een placed0 y a
contemporary taxonomist0 in the same species as their parents1 With hindsi!ht0 and for reasons that
are not far from le!alistic0 modern taxonomists insist on tyin! a lael around each fossil0 %hich
must say somethin! like A"stralopithec"s or )omo1 "useum laels are positively not allowed to
say (half%ay et%een A"stralopithec"s african"s and )omo habilis(1 )istory:deniers seiFe upon this
namin! convention as thou!h it %ere eidence of a lack of intermediates in the real %orld1 6ou
mi!ht as %ell say there is no such thin! as an adolescent ecause every sin!le person you look at
turns out to e either a votin! adult 7ei!hteen or over8 or a non:votin! child 7under ei!hteen81 #t(s
tantamount to sayin! that the le!al necessity for a votin! a!e threshold proves that adolescents don(t
Back to the fossils a!ain1 #f the creationist apolo!ists are ri!ht0 A"stralopithec"s is (Hust an
ape(0 so its o%n predecessors are irrelevant to the search for (missin! links(1 *evertheless0 %e may as
%ell look at them1 There are a fe%0 aleit rather fra!mentary0 traces1 Ardipithec"s0 %hich lived >:=
million years a!o0 is kno%n mainly from teeth0 ut enou!h cranial and foot ones have een found
to su!!est0 at least to most anatomists %ho have attended to it0 that it %alked upri!ht1 "uch the
same conclusion has een dra%n y the respective discoverers of t%o even older fossils0 !rrorin
7("illennium "an(8 and Sahelanthrop"s 7(Toumai(0 elo%81
Sahelanthrop"s is remarkale in ein! very old 7six million years0 close in a!e to the
common ancestor %ith chimpanFees8 and in ein! found far %est of the Rift +alley 7in Chad0 %here
its nickname0 (Toumai(0 means (hope of life(81 Other palaeoanthropolo!ists are sceptical of the claims
to ipedality that have een made on ehalf of !rrorin and Sahelanthrop"s y their discoverers1
&nd0 as a cynic mi!ht note0 for each such prolematic fossil some of the douters include the
discoverers of the othersK
Palaeoanthropolo!y0 more than other fields of science0 is notoriously pla!ued : or is it
enlivenedB : y rivalries1 We have to admit that the fossil record connectin! the upri!ht:%alkin! ape
A"stralopithec"s to the 7presumaly8 Euadrupedal ancestor that %e share %ith chimpanFees is still
poor1 We don(t kno% ho% our ancestors rose on to their hind le!s1 We need more fossils1 But let(s at
least reHoice in the !ood fossil record that %e : unlike 'ar%in : can enHoy0 sho%in! us the
evolutionary transition from A"stralopithec"s0 %ith its chimpanFee:siFed rain0 to modern )omo
sapiens %ith our alloon:like skull and i! rain1
Throu!hout this section # have reproduced pictures of skulls and encoura!ed you to compare
them1 6ou perhaps noticed0 for example0 the protrusion of the muFFle in some fossils0 or of the
ro% rid!e1 Sometimes the differences are Euite sutle0 %hich aids appreciation of the !radual
transitions from one fossil to a later one1 But no% # %ant to introduce a complication0 %hich %ill
develop into an interestin! point in its o%n ri!ht1 The chan!es that take place %ithin an individual(s
lifetime0 as it !ro%s up0 are in any case much more dramatic than the chan!es %e see as %e
compare adults in successive !enerations1
Chi"#an-ee shortly *e$ore *irth
The skull elo% elon!s to a chimpanFee shortly efore irth1 #t is oviously completely
different from the adult chimpanFee skull sho%n on pa!e 2;@ and far more like a human 7adult
human as %ell as ay human81 There(s a much:reproduced picture 7reproduced a!ain overleaf8 of
an infant and an adult chimpanFee0 %hich is often used to illustrate the interestin! idea that in
human evolution Huvenile characteristics %ere retained into adulthood 7or : %hich may not
necessarily e Euite the same thin! : %e ecome sexually mature %hen our odies are still Huvenile81
# thou!ht the picture looked too !ood to e true0 and # sent it to my collea!ue 'esmond "orris for
his expert opinion1 Could it e a fake0 # asked himB )ad he ever seen a youn! chimp lookin! Euite
so humanB 'r "orris is sceptical aout the ack and shoulders ut is happy aout the head itself1
(Chimps are characteristically hunched:up in posture and this one has a %onderfully upri!ht human
neck1 But if you Hust take the head alone0 the picture can e trusted1( Sheila -ee0 the pulishers(
picture researcher for this ook0 tracked do%n the ori!inal source of this famous photo!raph0 an
expedition to the Con!o in 2959:2= mounted y the &merican "useum of *atural )istory1 The
animals %ere dead %hen photo!raphed0 and she points out that the photo!rapher0 )erert -an!0 %as
also a taxidermist1 #t %ould e temptin! to surmise that the oddly human posture of the ay
chimpanFee is due to ad stuffin! : %ere it not for the fact that0 accordin! to the museum0 -an!
photo!raphed his specimens before stuffin! them1 *evertheless0 the posture of a dead chimp can e
adHusted in the %ay that the posture of a live chimp cannot1 'esmond "orris(s conclusion seems to
stand up1 The human:like posture of the ay chimpanFee(s shoulders may e suspect0 ut the head
is reliale1
1an''s #hotos o$ in$ant an% a%(lt hi"#an-ees
Takin! the head at face value0 even if the shoulders can(t Euite ear the urden of
authenticity0 you can immediately see ho% a comparison of adult fossil skulls mi!ht mislead us1 Or0
to put it more constructively0 the dramatic difference et%een adult and Huvenile heads sho%s us
ho% easily a characteristic such as muFFle protrusion mi!ht chan!e in Hust the ri!ht direction to
ecome more : or indeed less : human1 ChimpanFee emryolo!y (kno%s( ho% to make a human:like
head0 ecause it does it for every chimp as it passes throu!h its infant years1 #t seems hi!hly
plausile that0 as A"stralopithec"s evolved throu!h various intermediates to )omo sapiens0
shortenin! the muFFle all alon! the %ay0 it did so y the ovious route of retainin! Huvenile
characteristics into adulthood 7the process called neoteny0 mentioned in Chapter 381 #n any case0 a
!reat deal of evolutionary chan!e consists of chan!es in the rate at %hich certain parts !ro%0
relative to other parts1 This is called heterochronic 7(differently timed(8 !ro%th1 # suppose %hat #
%ant to say is that evolutionary chan!e is a doddle0 once you accept the oserved facts of
emryolo!ical chan!e1 Emryos are shaped y differential !ro%th : different its of them !ro% at
different rates1 & ay chimpanFee(s skull chan!es into an adult(s skull via relatively fast !ro%th of
the ones of the Ha%s and muFFle compared to other ones of the skull1 To repeat0 every animal of
every species chan!es0 durin! its o%n emryolo!ical development0 far more dramatically than the
typical adult form chan!es from !eneration to !eneration as the !eolo!ical a!es !o y1 &nd this is
my cue for a chapter on emryolo!y and its relevance to evolution1
L Predictaly0 the Pekin! fossil is no% sometimes called BeiHin! "an1 Why0 since %e are
talkin! En!lish rather than Chinese0 do %e !o alon! %ith (BeiHin!( at all0 %hen referrin! to China(s
capitalB There(s a rather charmin! pro!ramme on British television called Gr"mpy !ld %en0 %hich
is a !enially edited collection of !rouses and !riFFles of Hust this kind1 #f # %ere on it0 # %ould say
somethin! like the follo%in!1 We don(t da on a splash of Eau de Doln to dro%n out the smell of
"umai 'uck0 or !o %altFin! to the strains of (The Blue 'unaH( or (Tales from the Wien Woods(1
We don(t compare *eville Chamerlain0 the "an of "unchen0 to *apoleon(s retreat from "oskva1
*or yet 7thou!h !ive it time8 do %e take our snufflin! little pet BeiH for %alkies1 What(s %ron! %ith
Pekin!0 %hen it(s the En!lish lan!ua!e %e are speakin!B # %as deli!hted recently to meet a memer
of the British diplomatic corps0 fluent in "andarin0 %ho had played a leadin! role in our emassy in
%hat he insisted on callin! Pekin!1
L Pen name #sak 'inesen0 ut # like to use her real name ecause # spent my earliest
childhood near Daren0 the villa!e (at the foot of the *!on! )ills( %hich is still named after her1
L $nlike diseases0 %hich are often named after their discoverers0 ne% species are named by
their discoverers ut never after themselves1 #t is a nice opportunity for a iolo!ist to honour the
name of another0 or0 as in this case0 a enefactor1 *ot surprisin!ly0 my distin!uished collea!ue the
late W1 '1 )amilton %as several times honoured in this %ay1 &r!ualy one of 'ar%in(s !reatest
successors of the t%entieth century0 he had a lu!urious manner reminiscent of &1 &1 "ilne(s
Eeyore 7not the deplorale Walt 'isney version0 of course81 )amilton %as once on a small oat on
an expedition up the &maFon %hen he %as stun! y a %asp1 Dno%in! %hat a !reat entomolo!ist he
%as0 his companion said0 (Bill0 do you kno% the name of that %aspB( (6es0( Bill murmured !loomily
in his most Eeyoreish voice1 (&s a matter of fact it(s named after me1(
6O$ '#' #T 6O$RSE-, #* *#*E "O*T)S
T )&T irascile !enius .1 B1 S1 )aldane0 %ho did so much else esides ein! one of the
three leadin! architects of neo:'ar%inism0 %as once challen!ed y a lady after a pulic lecture1 #t(s
a %ord:of:mouth anecdote0 and .ohn "aynard Smith is sadly not availale to confirm the exact
%ords0 ut this is approximately ho% the exchan!e %entA
Eol"tion scepticA Professor )aldane0 even !iven the illions of years that you say %ere
availale for evolution0 # simply cannot elieve it is possile to !o from a sin!le cell to a
complicated human ody0 %ith its trillions of cells or!aniFed into ones and muscles and nerves0 a
heart that pumps %ithout ceasin! for decades0 miles and miles of lood vessels and kidney tuules0
and a rain capale of thinkin! and talkin! and feelin!1
-BSA But madam0 you did it yourself1 &nd it only took you nine months1
The Euestioner %as perhaps momentarily thro%n off alance y the veerin! unexpectedness
of )aldane(s reply1 Wind taken out of sails %ould have seemed an understatement1 But maye in
one respect )aldane(s retort left her unsatisfied1 # don(t kno% %hether she asked a supplementary
ut0 if so0 it mi!ht have !one alon! these linesA
Eol"tion scepticA &h yes0 ut the developin! emryo follo%s !enetic instructions1 #t is the
instr"ctions for ho% to uild a complicated ody that you0 Professor )aldane0 claim evolved y
natural selection1 &nd # still find it hard to elieve0 even !iven a illion years for that evolution1
Perhaps she had a point1 &nd even if a divine intelli!ence did prove to e ultimately
responsile for desi!nin! livin! complexity0 it is definitely not true that he fashions livin! odies in
anythin! like the %ay that clay modellers0 for example0 or carpenters0 potters0 tailors or car
manufacturers !o aout their tasks1 We may e (%onderfully developed( ut %e are not (%onderfully
made'1 When children sin!0 ()e made their !lo%in! colours G )e made their tiny %in!s(0 L they are
utterin! a childishly ovious falsehood1 Whatever else God does0 he certainly doesn(t make !lo%in!
colours and tiny %in!s1 #f he did anythin! at all0 it %ould e to supervise the emryonic
development of thin!s0 for example y splicin! to!ether seEuences of !enes that direct a process of
automated development1 Win!s are not made0 they !ro% : pro!ressively : from lim uds inside an
God0 to repeat this important point0 %hich ou!ht to e ovious ut isn(t0 never made a tiny
%in! in his eternal life1 #f he made anythin! 7he didn(t in my vie%0 ut let it pass0 that(s not %hat #(m
aout here80 %hat he made %as an emryolo!ical recipe0 or somethin! like a computer pro!ram for
controllin! the emryonic development of a tiny %in! 7plus lots of other thin!s too81 Of course0
God mi!ht claim that it is Hust as clever0 Hust as reathtakin! a feat of skill0 to desi!n a recipe or a
pro!ram for a %in!0 as to make a %in!1 But for the moment0 # Hust %ant to develop the distinction
et%een makin$ somethin! like a %in!0 and %hat really happens in emryolo!y1
The early history of emryolo!y %as riven et%een t%o opposin! doctrines called
preformationism and epi!enesis1 The distinction et%een them is not al%ays clearly understood0 so
# shall spend a little time explainin! these t%o terms1 The preformationists elieved that the e!! 7or
sperm0 for the preformationists %ere sudivided into (ovists( versus (spermists(8 contained a tiny
miniature ay or (homunculus(1 &ll the parts of the ay %ere intricately in place0 correctly
disposed to each other0 %aitin! only to e inflated like a compartmentaliFed alloon1 This raises
ovious prolems1 ,irst0 at least in its early naive form0 it reEuires %hat everyody kno%s to e
falseA that %e inherit only from one parent : the mother for the ovists0 the father for the spermists1
Second0 preformationists of this kind had to face a Russian:doll:style infinite re!ress of homunculi
%ithin homunculi : or if not infinite0 at least lon! enou!h to take us ack to Eve 7&dam for the
spermists81 The only escape from the re!ress %ould e to construct the homunculus afresh in every
!eneration y elaorately scannin! the adult ody of the previous !eneration1 This (inheritance of
acEuired characteristics( doesn(t happen : other%ise .e%ish oys %ould e orn %ithout foreskins0
and !ym:freEuentin! ody:uilders 7ut not their couch:potato t%ins8 %ould conceive aies %ith
ripplin! six:packs0 pecs and !lutes1
To e fair to the preformationists0 they did face up0 fairly and sEuarely0 to the lo!ical
necessity of the re!ress0 ho%ever asurd it seemed1 &t least some of them really did elieve that the
first %oman 7or man8 contained miniaturiFed emryos of all her descendants0 nested inside each
other like Russian dolls1 &nd there is a sense in %hich they had to elieve thatA a sense that is %orth
mentionin! ecause it prefi!ures the nu of this chapter1 #f you elieve &dam %as (made( rather than
ein! orn0 you imply that &dam didn(t have !enes : or at least didn(t need them in order to
develop1 &dam had no emryolo!y ut Hust spran! into existence1 & related inference led the
+ictorian %riter Philip Gosse 7the father in Edmund Gosse(s ,ather and Son8 to %rite a ook called
!mphalos 7Greek for (navel(8 ar!uin! that &dam must have had a navel0 even thou!h he %as never
orn1 & more sophisticated conseEuence of omphalo!ical reasonin! %ould e that stars %hose
distance from us is more than a fe% thousand li!ht years must have een created %ith ready:made
li!ht eams stretchin! almost all the %ay to us : other%ise %e %ouldn(t e ale to see them until the
distant futureK "akin! fun of omphalo!y sounds frivolous0 ut there is a serious point here aout
emryolo!y0 %hich is the suHect of this chapter1 #t is Euite a difficult point to !rasp : indeed0 # am
only in the process of !raspin! it myself : and # am approachin! it from various directions1
,or the reasons !iven0 preformationism0 at least in its ori!inal (Russian doll( version0 %as
al%ays a non:starter1 #s there a version of preformationism that could e sensily revived in the
'*& a!eB Well0 perhaps0 ut # dout it1 Textooks of iolo!y repeat time and a!ain that '*& is a
(lueprint( for uildin! a ody1 #t isn(t1 & true lueprint of0 say0 a car or a house emodies a one:to:
one mappin! from paper to finished product1 #t follo%s from this that a lueprint is reversile1 #t is
as easy to !o from house to lueprint as the other %ay around0 precisely ecause it is a one:to:one
mappin!1 &ctually0 it(s easier0 ecause you have to b"ild the house0 ut you only have to take some
measurements and then draw the lueprint1 #f you take an animal(s ody0 no matter ho% many
detailed measurements you take0 you can(t reconstruct its '*&1 That(s %hat makes it false to say
that '*& is a lueprint1
#t is theoretically possile to ima!ine : maye that(s the %ay thin!s %ork on some alien
planet : that '*& mi!ht have een a coded description of a odyA a kind of three:dimensional map
rendered into the linear code of '*& (letters(1 That really %ould e reversile1 Scannin! the ody to
make a !enetic lueprint is not a totally ridiculous idea1 #f that is ho% '*& %orked0 %e could
represent it as a kind of neo:preformationism1 #t %ouldn(t raise the spectre of the Russian dolls1 #t
isn(t clear to me %hether it %ould raise the spectre of inheritance from only one parent1 '*& has a
reathtakin!ly precise %ay of intersplicin! half the paternal information %ith exactly half the
maternal information0 ut ho% %ould it !o aout intersplicin! half a scan of the mother(s ody %ith
half a scan of the father(s odyB -et it passA this is all so far from reality1
'*&0 then0 is emphatically not a lueprint1 $nlike &dam0 %ho %as fashioned directly into
his adult form0 all real odies develop and !ro% from a sin!le cell throu!h the intermediate sta!es
of emryo0 foetus0 ay0 child and adolescent1 "aye in some alien %orld livin! creatures assemle
themselves from tip to toe as an ordered set of three:dimensional io:pixels read out from a coded
scan line1 But that is not the %ay thin!s %ork on our planet0 and actually # think there are reasons :
%hich # have dealt %ith else%here and so %on(t !o into here : %hy it could never e so on any
The historical alternative to preformationism is epi!enesis1 #f preformationism is all aout
lueprints0 epi!enesis is aout somethin! more like a recipe or a computer pro!ram1 The Shorter
!xford En$lish Dictionary(s definition is rather modern0 and #(m not sure that &ristotle0 %ho coined
the %ord0 %ould reco!niFe itA
e#i'enesisA & theory of the development of an or!anism y pro!ressive differentiation of an
initially undifferentiated %hole1 L
Principles of Deelopment0 y -e%is Wolpert and collea!ues0 descries epi!enesis as the
idea that ne% structures arise pro!ressively1 There is a sense in %hich epi!enesis is self:evidently
true0 ut details matter and the devil is in the cliche1 )o% does the or!anism develop pro!ressivelyB
)o% does an initially undifferentiated %hole (kno%( ho% to differentiate pro!ressively0 if not y
follo%in! a lueprintB The distinction # %ant to make in this chapter0 %hich lar!ely corresponds to
the distinction et%een preformationism and epi!enesis0 is the distinction et%een planned
architecture and self0assembly1 The meanin! of planned architecture is clear to us ecause %e see it
all around us in our uildin!s and other artefacts1 Self:assemly is less familiar0 and %ill need some
attention from me1 #n the field of development0 self:assemly occupies a position analo!ous to
natural selection in evolution0 althou!h it is definitely not the same process1 Both achieve0 y
automatic0 non:delierate0 unplanned means0 results that look0 to a superficial !aFe0 as thou!h they
%ere meticulously planned1
.1 B1 S1 )aldane spoke simple truth to his sceptical Euestioner0 ut he %ould not have denied
that there is mystery0 ver!in! on the miraculous 7ut never Euite !ettin! there8 in the very fact that a
sin!le cell !ives rise to a human ody in all its complexity1 &nd the mystery is only some%hat
miti!ated y the feat(s ein! achieved %ith the aid of '*& instructions1 The reason the mystery
remains is that %e find it hard to ima!ine0 even in principle0 ho% %e mi!ht set aout %ritin! the
instructions for uildin! a ody in the %ay that the ody is in fact uilt0 namely y %hat # have Hust
called (self:assemly(0 %hich is related to %hat computer pro!rammers sometimes call a (ottom:up(0
as opposed to (top:do%n(0 procedure1
&n architect desi!ns a !reat cathedral1 Then0 throu!h a hierarchical chain of command0 the
uildin! operation is roken do%n into separate departments0 %hich reak it do%n further into su:
departments0 and so on until instructions are finally handed out to individual masons0 carpenters and
!laFiers0 %ho !o to %ork until the cathedral is uilt0 lookin! pretty much like the architect(s ori!inal
dra%in!1 That(s top:do%n desi!n1
Bottom:up desi!n %orks completely differently1 # never elieved this0 ut there used to e a
myth that some of the finest medieval cathedrals in Europe had no architect1 *oody desi!ned the
cathedral1 Each mason and carpenter %ould usy himself0 in his o%n skilled %ay0 %ith his o%n little
corner of the uildin!0 payin! scant attention to %hat the others %ere doin! and no attention to any
overall plan1 Someho%0 out of such anarchy0 a cathedral %ould emer!e1 #f that really happened0 it
%ould e ottom:up architecture1 *ot%ithstandin! the myth0 it surely didn(t happen like that for
cathedrals1L But that pretty much is %hat happens in the uildin! of a termite mound or an ant(s nest
: and in the development of an emryo1 #t is %hat makes emryolo!y so remarkaly different from
anythin! %e humans are familiar %ith0 in the %ay of construction or manufacture1
The same principle %orks for certain types of computer pro!ram0 for certain types of animal
ehaviour0 and : rin!in! the t%o to!ether : for computer pro!rams that are desi!ned to simulate
animal ehaviour1 Suppose %e %anted to understand the flockin! ehaviour of starlin!s1 There are
some stunnin! films availale on 6ouTue0 from %hich the stills on colour pa!e 2< are taken1
These alletic manoeuvres %ere photo!raphed over Otmoor0 near Oxford0 y 'ylan Winter1 What is
remarkale aout the starlin!s( ehaviour is that0 despite all appearances0 there is no choreo!rapher
and0 as far as %e kno%0 no leader1 Each individual ird is Hust follo%in! local rules1
The numers of individual irds in these flocks can run into thousands0 yet they almost
literally never collide1 That is Hust as %ell for0 !iven the speed at %hich they fly0 any such impact
%ould severely inHure them1 Often the %hole flock seems to ehave as a sin!le individual0 %heelin!
and turnin! as one1 #t can look as thou!h the separate flocks are movin! throu!h each other in
opposite directions0 maintainin! their coherence intact as separate flocks1 This makes it seem almost
miraculous0 ut actually the flocks are at different distances from the camera and do not literally
move throu!h each other1 #t adds to the aesthetic pleasure that the ed!es of the flocks are so sharply
defined1 They don(t peter off !radually0 ut come to an arupt oundary1 The density of the irds
Hust inside the oundary is no less than in the middle of the flock0 %hile it is Fero outside the
oundary1 &s soon as you think aout it in that %ay0 isn(t it %ondrously surprisin!B
The %hole performance %ould make a more than usually ele!ant screensaver on a computer1
6ou %ouldn(t %ant a real film of starlin!s ecause your screensaver %ould repeat the same identical
alletic moves over and over0 and therefore %ouldn(t exercise all the pixels eEually1 What you
%ould %ant is a computer sim"lation of starlin! flocksJ and0 as any pro!rammer %ill tell you0
there(s a ri!ht %ay and a %ron! %ay to do it1 'on(t try to choreo!raph the %hole allet : that %ould
e terrily ad pro!rammin! style for this kind of task1 # need to talk aout the etter %ay to do it
ecause somethin! like it is almost certainly ho% the irds themselves are pro!rammed0 in their
rains1 "ore to the point0 it is a !reat analo!y for ho% emryolo!y %orks1
)ere(s ho% to pro!ram flockin! ehaviour in starlin!s1 'evote almost all your effort to
pro!rammin! the ehaviour of a sin!le individual ird1 Build into your roo:starlin! detailed rules
for ho% to fly0 and ho% to react to the presence of nei!hourin! starlin!s0 dependin! on their
distance and relative position1 Build in rules for ho% much %ei!ht to !ive to the ehaviour of
nei!hours0 and ho% much %ei!ht to !ive to individual initiative in chan!in! direction1 These
model rules %ould e informed y careful measurements of real irds in action1 Endo% your
cyerird %ith a certain tendency to vary its rules at random1 )avin! %ritten a complicated pro!ram
to specify the ehavioural rules of a sin!le starlin!0 no% comes the definitive step that # am
emphasiFin! in this chapter1 Don't try to pro!ram the ehaviour of a %hole flock0 as an earlier
!eneration of computer pro!rammers mi!ht have done1 #nstead0 clone the sin!le computer starlin!
you have pro!rammed1 "ake a thousand copies of your roo:ird0 maye all the same as each
other0 or maye %ith some sli!ht random variation amon! them in their rules1 &nd no% (release(
thousands of model starlin!s in your computer0 so they are free to interact %ith each other0 all
oeyin! the same rules1
#f you(ve !ot the ehavioural rules ri!ht for a sin!le starlin!0 a thousand computer starlin!s0
each one a dot on the screen0 %ill ehave like real starlin!s flockin! in %inter1 #f the flockin!
ehaviour isn(t Euite ri!ht0 you can !o ack and adHust the ehaviour of the individual starlin!0
perhaps in the li!ht of further measurements of the ehaviour of real starlin!s1 *o% clone up the
ne% version a thousand times0 in place of the thousand that didn(t Euite %ork1 Deep iteratin! your
repro!rammin! of the cloned:up sin!le starlin!0 until the flockin! ehaviour of thousands of them
on the screen is a satisfyin!ly realistic screensaver1 Callin! it (Boids(0 Crai! Reynolds %rote a
pro!ram alon! these lines 7not specifically for starlin!s8 in 29;<1
The key point is that there is no choreo!rapher and no leader1 Order0 or!aniFation0 structure
: these all emer$e as y:products of rules %hich are oeyed locally and many times over0 not
!loally1 &nd that is ho% emryolo!y %orks1 #t is all done y local rules0 at various levels ut
especially the level of the sin!le cell1 *o choreo!rapher1 *o conductor of the orchestra1 *o central
plannin!1 *o architect1 #n the field of development0 or manufacture0 the eEuivalent of this kind of
pro!rammin! is self0assembly1
The ody of a human0 an ea!le0 a mole0 a dolphin0 a cheetah0 a leopard fro!0 a s%allo%A
these are so eautifully put to!ether0 it seems impossile to elieve that the !enes that pro!ram their
development don(t function as a lueprint0 a desi!n0 a master plan1 But noA as %ith the computer
starlin!s0 it is all done y individual cells oeyin! local rules1 The eautifully (desi!ned( ody
emer$es as a conseEuence of rules ein! locally oeyed y individual cells0 %ith no reference to
anythin! that could e called an overall !loal plan1 The cells of a developin! emryo %heel and
dance around each other like starlin!s in !i!antic flocks1 There are differences0 and they are
important1 $nlike starlin!s0 cells are physically attached to each other in sheets and locksA their
(flocks( are called (tissues(1 When they %heel and dance like miniature starlin!s0 the conseEuence is
that three:dimensional shapes are formed0 as tissues inva!inate in response to the movements of
cellsJ L or s%ell or shrink due to local patterns of !ro%th and cell death1 The analo!y # like for this
is the paper:foldin! art of ori!ami0 su!!ested y the distin!uished emryolo!ist -e%is Wolpert in
his ook The Tri"mph of the EmbryoJ ut efore comin! to that # need to clear out of the %ay some
alternative analo!ies that mi!ht come to mind : analo!ies from amon! human crafts and
manufacturin! processes1
&*&-OG#ES ,OR 'E+E-OP"E*T
#t is surprisin!ly hard to find a !ood analo!y for the development of livin! tissue0 ut you
can find partial similarities to particular aspects of the process1 & recipe captures somethin! of the
truth0 and it is an analo!y that # sometimes use0 to explain %hy (lueprint( is not appropriate1 $nlike
a lueprint0 a recipe is irreversile1 #f you follo% a cake recipe step y step0 you(ll end up %ith a
cake1 But you can(t take a cake and reconstruct the recipe : certainly not the exact %ords of the
recipe : %hereas0 as %e have seen0 you could take a house and reconstruct somethin! close to the
ori!inal lueprint1 This is ecause of the one:to:one mappin! et%een its of house and its of
lueprint1 With conspicuous exceptions such as the cherry on top0 there is no one:to:one mappin!
et%een its of cake and the %ords0 say0 or sentences of its recipe1
What other analo!ies to human manufacturin! mi!ht there eB Sculpture is mostly %ay off
the mark1 & sculptor starts %ith a chunk of stone or %ood and fashions it y sutraction0 chippin!
a%ay until the desired shape is all that remains1 There is0 admittedly0 a some%hat sharp resemlance
to one particular process in emryolo!y called apoptosis1 &poptosis is pro!rammed cell death0 and
it is involved0 for example0 in the development of fin!ers and toes1 #n the human emryo0 the
fin!ers and toes are all Hoined1 #n the %om0 you and # had %eed feet and hands1 The %ein!
disappeared 7in most peopleA there are occasional exceptions8 throu!h pro!rammed cell death1 That
is a it reminiscent of the %ay a sculptor carves out a shape0 ut it is not common enou!h or
important enou!h to capture ho% emryolo!y normally %orks1 Emryolo!ists may riefly think
(sculptor(s chisel(0 ut they don(t let the thou!ht lin!er for lon!1
Some sculptors %ork not y sutractive carvin! ut y takin! a lump of clay0 or soft %ax0
and kneadin! it into shape 7%hich may suseEuently e cast0 in ronFe for example81 That a!ain is
not a !ood analo!y for emryolo!y1 *or is the craft of tailorin! or dressmakin!1 Pre:existin! cloth
is cut0 to shapes set out in a pre:planned pattern0 then se%n to!ether %ith other cut:out shapes1 They
are often then turned inside out to dis!uise the seams : and that it0 at least0 is a !ood analo!y to
certain parts of emryolo!y1 But in !eneral0 emryolo!y is no more like tailorin! than it is like
sculpture1 Dnittin! mi!ht e etter0 in that the %hole shape of a s%eater0 say0 is uilt up from
numerous individual stitches0 like individual cells1 But there are etter analo!ies0 as %e shall see1
)o% aout the assemly of a car0 or other complicated machine0 on a factory assemly lineA
is that a !ood analo!yB -ike sculpture and tailorin!0 assemly of pre:faricated parts is an efficient
%ay to make somethin!1 #n a car factory0 parts are pre:made0 often y castin! in moulds in a
foundry 7and there is0 # think0 nothin! remotely like castin! in emryolo!y81 Then the pre:made
parts are rou!ht to!ether on an assemly line and scre%ed0 riveted0 %elded or !lued to!ether0 step
y step accordin! to a precisely dra%n plan1 Once a!ain0 emryolo!y has nothin! resemlin! a
previously dra%n plan1 But there are resemlances to the ordered stickin! to!ether of pre:assemled
parts0 as %hen0 in a car assemly plant0 previously manufactured carurettors and distriutor heads
and fan elts and cylinder heads are rou!ht to!ether and Hoined in correct apposition1
Belo% are three kinds of virus1 On the left is the toacco mosaic virus 7T"+80 %hich
parasitiFes toacco plants and other memers of the family Solanaceae0 such as tomatoes1 #n the
middle is an adenovirus0 %hich infects the respiratory system in many animals0 includin! us1 On the
ri!ht is the T> acteriopha!e0 %hich parasitiFes acteria1 #t looks like a lunar lander0 and it ehaves
rather like one0 (landin!( on the surface of a acterium 7%hich is very much lar!er8 then lo%erin!
itself on its spidery (le!s(0 then thrustin! a proe do%n the middle0 throu!h the acterium(s cell %all0
and inHectin! its '*& inside1 The viral '*& then hiHacks the protein:makin! machinery of the
acterium0 %hich is suverted into makin! ne% viruses1 The other t%o viruses in the picture do
somethin! similar0 althou!h they don(t look or ehave like lunar landers1 #n all cases their !enetic
material hiHacks the protein:makin! apparatus of the host cell and diverts its molecular production
line to churnin! out viruses instead of its normal products1
Three &in%s o$ .ir(s
"ost of %hat you see in the pictures is the protein container for the !enetic material0 and in
7(lunar lander(8 T>(s case the machinery for infectin! the host1 What is interestin! is the %ay in
%hich this protein apparatus is put to!ether1 #t really is self:assemled1 Each virus is assemled
from several previously made protein molecules1 Each protein molecule0 in a %ay that %e shall see0
has previously self:assemled into a characteristic (tertiary structure( under the la%s of chemistry
!iven its particular seEuence of amino acids1 &nd then0 in the virus0 the protein molecules Hoin up
%ith each other to form a so:called (Euaternary structure(0 a!ain y follo%in! local rules1 There is no
!loal plan0 no lueprint1
The protein su:units0 %hich link up like -e!o ricks to form the Euaternary structure0 are
called capsomeres1 *otice ho% !eometrically perfect these little constructions are1 The adenovirus
in the middle has exactly 3=3 capsomeres0 dra%n here as little alls0 arran!ed in an icosahedron1
The icosahedron is that Platonic perfect solid that has 35 trian!ular faces1 The capsomeres are
arran!ed into an icosahedron not y any kind of master plan or lueprint ut simply y each one of
them oeyin! the la%s of chemical attraction locally %hen it umps into others like itself1 This is
ho% crystals are formed0 and0 indeed0 the adenovirus could e descried as a very small hollo%
crystal1 The (crystalliFation( of viruses is an especially eautiful example of the (self:assemly( that #
am toutin! as a maHor principle y %hich livin! creatures are put to!ether1 The T> (lunar lander(
pha!e also has an icosahedron for its main '*& receptacle0 ut its self:assemled Euaternary
structure is more complex0 incorporatin! additional protein units0 assemled accordin! to different
local rules0 in the inHection apparatus and the (le!s( that are attached to the icosahedron1
Returnin! from viruses to the emryolo!y of lar!er creatures0 # come to my favourite
analo!y amon! human construction techniEuesA ori!ami1 Ori!ami is the art of constructive paper:
foldin!0 developed to its most advanced level in .apan1 The only ori!ami creation # kno% ho% to
make is the (Chinese .unk(1 # %as tau!ht it y my father0 %ho learned it in a craFe that s%ept throu!h
his oardin! school durin! the 2935s1L One iolo!ically realistic feature is that the (emryolo!y( of
the Chinese Hunk passes throu!h several intermediate (larval( sta!es0 %hich are in themselves
pleasin! creations0 Hust as a caterpillar is a eautiful0 %orkin! intermediate on the %ay to a utterfly0
%hich it scarcely resemles at all1 Startin! %ith a simple sEuare piece of paper0 and simply foldin!
it : never cuttin! it0 never !luein! it and never importin! any other pieces : the procedure takes us
throu!h three reco!niFale (larval sta!es(A a (catamaran(0 a (ox %ith t%o lids( and a (picture in a
frame(0 efore culminatin! in the (adult( Chinese Hunk itself1 #n favour of the ori!ami analo!y0 %hen
you first are tau!ht ho% to make a Chinese Hunk0 not only the Hunk itself ut each of the three (larval(
sta!es : catamaran0 cupoard0 picture frame : comes as a surprise1 6our hands may do the foldin!0
ut you are emphatically not follo%in! a lueprint for a Chinese Hunk0 or for any of the larval
sta!es1 6ou are follo%in! a set of foldin! rules that seem to have no connection %ith the end
product0 until it finally emer!es like a utterfly from its chrysalis1 So the ori!ami analo!y captures
somethin! of the importance of (local rules( as opposed to a !loal plan1
&lso in favour of the ori!ami analo!y0 foldin!0 inva!ination and turnin! inside out are some
of the favourite tricks used y emryonic tissues %hen makin! a ody1 The analo!y %orks
especially %ell for the early emryonic sta!es1 But it has its shortcomin!s0 and here are t%o ovious
ones1 ,irst0 human hands are needed to do the foldin!1 Second0 the developin! paper (emryo(
doesn(t !ro% lar!er1 #t ends up %ei!hin! exactly as much as %hen it started1 To ackno%led!e the
difference0 # shall sometimes refer to iolo!ical emryolo!y as (inflatin! ori!ami(0 rather than Hust
Chinese =(n& *y ori'a"i, ,ith three ' sta'es': 'ata"aran', '*o/ ,ith t,o li%s'
an% '#it(re in a $ra"e'
&ctually0 these t%o shortcomin!s kind of cancel each other out1 The sheets of tissue that
fold0 inva!inate and turn inside out in a developin! emryo do indeed !ro%0 and it is that very
!ro%th that provides part of the motive force %hich0 in ori!ami0 is supplied y the human hand1 #f
you %anted to make an ori!ami model %ith a sheet of livin! tissue instead of dead paper0 there is at
least a sportin! chance that0 if the sheet %ere to !ro% in Hust the ri!ht %ay0 not uniformly ut faster
in some parts of the sheet than in others0 this mi!ht automatically cause the sheet to assume a
certain shape : and even fold or inva!inate or turn inside out in a certain %ay : %ithout the need for
hands to do the stretchin! and foldin!0 and %ithout the need for any !loal plan0 ut only local
rules1 &nd actually it(s more than Hust a sportin! chance0 ecause it really happens1 -et(s call it (auto:
ori!ami(1 )o% does auto:ori!ami %ork in practice0 in emryolo!yB #t %orks ecause %hat happens
in the real emryo0 %hen a sheet of tissue !ro%s0 is that cells divide1 &nd differential !ro%th of the
different parts of the sheet of tissue is achieved y the cells0 in each part of the sheet0 dividin! at a
rate determined y local rules1 So0 y a roundaout route0 %e return to the fundamental importance
of ottom:up local rules as opposed to top:do%n !loal rules1 #t is a %hole series of 7far more
complicated8 versions of this simple principle that actually !o on in the early sta!es of emryonic
)ere(s ho% the ori!ami !oes in the early sta!es of verterate development1 The sin!le
fertiliFed e!! cell divides to make t%o cells1 Then the t%o divide to make four1 &nd so on0 %ith the
numer of cells rapidly doulin! and redoulin!1 &t this sta!e there is no !ro%th0 no inflation1 The
ori!inal volume of the fertiliFed e!! is literally divided0 as in slicin! a cake0 and %e end up %ith a
spherical all of cells %hich is the same siFe as the ori!inal e!!1 #t(s not a solid all ut a hollo%
one0 and it is called the lastula1 The next sta!e0 !astrulation0 is the suHect of a famous bon mot y
-e%is WolpertA (#t is not irth0 marria!e0 or death0 ut !astrulation0 %hich is truly the most
important time in your life1(
Gastrulation is a kind of microcosmic earthEuake %hich s%eeps over the lastula(s surface
and revolutioniFes its entire form1 The tissues of the emryo ecome massively reor!aniFed1
Gastrulation typically involves a dentin! of the hollo% all that is the lastula0 so that it ecomes
t%o:layered %ith an openin! to the outside %orld 7see the computer simulation on p1 34281 The
outer layer of this (!astrula( is called the ectoderm0 the inner layer is the endoderm0 and there are
also some cells thro%n into the space et%een the ectoderm and endoderm0 %hich are called
mesoderm1 Each of these three primordial layers is destined to make maHor parts of the ody1 ,or
example0 the outer skin and nervous system come from the ectodermJ the !uts and other internal
or!ans come from the endodermJ and the mesoderm furnishes muscle and one1
The next sta!e in the emryo(s ori!ami is called neurulation1 The dia!ram on the ri!ht sho%s
a cross:section throu!h the middle of the ack of a neurulatin! amphiian emryo 7it could e either
a fro! or a salamander81 The lack circle is the (notochord(0 a stiffenin! rod that acts as a precursor
of the ackone1 The notochord is dia!nostic of the phylum Chordata0 to %hich %e and all
verterates elon! 7althou!h %e0 like most modern verterates0 have it only %hen %e are emryos81
#n neurulation0 as in !astrulation0 inva!ination is much in evidence1 6ou rememer # said that the
nervous system comes from ectoderm1 Well0 here(s ho%1 & section of ectoderm inva!inates
7pro!ressively ack%ards alon! the ody like a Fip fastener80 rolls itself up into a tue0 and is
pinched off %here the sides of the tue (Fip up( so that it ends up runnin! the len!th of the ody
et%een the outer layer and the notochord1 That tue is destined to ecome the spinal cord0 the main
nerve trunk of the ody1 The front end of it s%ells up and ecomes the rain1 &nd all the rest of the
nerves are derived0 y suseEuent cell divisions0 from this primordial tue1 L
# don(t %ant to !et into the details of either !astrulation or neurulation0 except to say that
they are %onderful0 and that the metaphor of ori!ami holds up pretty %ell for oth of them1 # am
concerned %ith the !eneral principles y %hich emryos ecome more complicated throu!h
inflatin! ori!ami1 Belo% is one of the thin!s that sheets of cells are oserved to do durin! the course
of emryonic development0 for example durin! !astrulation1 6ou can easily see ho% this
inva!ination could e a useful move in inflatin! ori!ami0 and it does indeed play a maHor role in
oth !astrulation and neurulation1
In.a'ination in a sheet o$ ells
Gastrulation and neurulation are accomplished early in development and they affect the
%hole shape of the emryo1 #nva!ination and other (inflatin! ori!ami( manoeuvres achieve these
sta!es of early emryolo!y0 and they and similar tricks are involved later in development0 %hen
specialiFed or!ans like eyes and the heart are made1 But0 !iven that there are no hands to do the
foldin!0 y %hat mechanical process are these dynamic movements achievedB Partly0 as # have
already said0 y simple expansion itself1 Cells multiply all throu!h a sheet of tissue1 #ts area
therefore increases and0 havin! no%here else to !o0 it has little choice ut to uckle or inva!inate1
But the process is more controlled than that0 and it has een deciphered y a !roup of scientists
associated %ith the rilliant mathematical iolo!ist Geor!e Oster0 of the $niversity of California at
"O'E--#*G CE--S -#DE ST&R-#*GS
Oster and his collea!ues follo%ed the same strate!y %e considered earlier in this chapter for
a computer simulation of starlin!s flockin!1 #nstead of pro!rammin! the ehaviour of a %hole
lastula0 they pro!rammed a sin!le cell1 Then they (cloned up( lots of cells0 all the same0 and
%atched to see %hat happened %hen those cells !ot to!ether in the computer1 When # say they
pro!rammed the ehaviour of a sin!le cell0 it %ould e etter to say they pro!rammed a
mathematical model of a sin!le cell0 uildin! into the model certain kno%n facts aout a sin!le cell0
ut in simplified form1 Specifically0 it is kno%n that the interiors of cells are criss:crossed y
microfilamentsA sort of miniature elastic ands0 ut %ith the additional property that they are
capale of active contraction0 like t%itchin! muscle fires1 #ndeed0 the microfilaments use the same
principle of contraction as muscle fires1L The Oster model simplified the cell do%n to t%o
dimensions for dra%in! on a computer screen0 and %ith only half a doFen filaments0 strate!ically
placed in the cell0 as you see in the dia!ram aove1 #n the computer model0 all the microfilaments
%ere !iven certain Euantitative properties %ith names that mean somethin! to physicistsA a (viscous
dampin! coefficient( and an (elastic sprin! constant(1 *ever mind exactly %hat these meanA they are
the kinds of thin!s physicists like to measure in a sprin!1 &lthou!h it is proale that in a real cell
many filaments %ould e capale of contraction0 Oster and his collea!ues simplified matters y
endo%in! only one of their six filaments %ith this capacity1 #f they could !et realistic results even
after thro%in! a%ay some of the kno%n properties of a cell0 it %ould presumaly e possile to !et
at least as !ood results %ith a more complicated model that kept those properties in1 Rather than
allo%in! the one contractile filament in their model to contract at %ill0 they uilt into it a property
%hich is common in certain kinds of muscle fireA %hen stretched eyond a certain critical len!th0
the fire %ould respond y contractin! to a much shorter len!th than the normal eEuilirium len!th1
4iro$ila"ents insi%e 8ster's "o%el ell
So0 %e have our model of a sin!le cellA a !reatly simplified model consistin! of a t%o:
dimensional outline in %hich are strun! six elastic sprin!s0 one of %hich has the special property of
respondin! to an externally imposed stretch y actively contractin!1 That is sta!e one of the
modellin! process1 #n sta!e t%o0 Oster and his collea!ues cloned up a fe% doFen of their model
cells and arran!ed them in a circle0 like a 7t%o:dimensional8 lastula1 Then they took one cell and
t%eaked its contractile filament to provoke it into contractin!1 What happened next is almost too
%onderful to ear1 The model lastula !astrulatedK )ere are six screenshots sho%in! %hat happened
7a to f elo%81 & %ave of contraction spread side%ays from the cell that %as provoked0 and the all
of cells spontaneously inva!inated1
8ster's "o%el *last(la 'astr(latin'
#t !ets even etter1 Oster and his collea!ues tried the experiment0 on their computer model0
of lo%erin! the (firin! threshold( of the contractile filaments1 The result %as a %ave of inva!ination
that %ent further0 and actually pinched off a (neural tue( 7screenshots a to h0 overleaf81 #t is
important to understand %hat a model such as this really is1 #t is not an accurate representation of
neurulation1 Puite apart from the fact that it is t%o:dimensional and simplified in many other %ays0
the all of cells that (neurulated( 7screenshot a8 %as not a t%o:layered (!astrula( as it should have
een1 #t %as the same lastula:like startin! point as %e had for the model of !astrulation aove1 #t
doesn(t matterA models are not supposed to e totally accurate in every detail1 The model still sho%s
ho% easy it is to mimic various aspects of the ehaviour of cells in an early emryo1 The fact that
the t%o:dimensional (all( of cells responded spontaneously to the stimulus even thou!h the model
is simpler than the real situation makes this a more po%erful piece of evidence1 #t reassures us that
the evolution of the various procedures of early emryonic development need not have een all that
difficult1 *ote that it is the model that is simple0 not the phenomenon that it demonstrates1 That is
the hallmark of a !ood scientific model1
For"ation o$ 'ne(ral t(*e' in 8ster's "o%el
"y purpose in expoundin! the Oster models has een to sho% the !eneral kind of principle
y %hich sin!le cells can interact %ith each other to uild a ody0 %ithout any lueprint
representin! the %hole ody1 Ori!ami:like foldin!0 Oster:style inva!ination and pinchin! offA these
are Hust some of the simplest tricks for uildin! emryos1 Other more elaorate ones come into play
later in emryonic development1 ,or example0 in!enious experiments have sho%n that nerve cells0
%hen they !ro% out from the spinal cord0 or from the rain0 find their %ay to their end or!an not y
follo%in! any kind of overall plan ut y chemical attraction0 rather as a do! sniffs around to find a
itch in season1 &n early classic experiment y the *oel PriFe:%innin! emryolo!ist Ro!er Sperry
illustrates the principle perfectly1 Sperry and a collea!ue took a tadpole and removed a tiny sEuare
of skin from the ack1 They removed another sEuare0 the same siFe0 from the elly1 They then
re!rafted the t%o sEuares0 ut each in the other(s placeA the elly skin %as !rafted on the ack0 and
the ack skin on the elly1 When the tadpole !re% up into a fro!0 the result %as rather pretty0 as
experiments in emryolo!y often areA there %as a neat posta!e stamp of %hite elly skin in the
middle of the dark0 mottled ack0 and another neat posta!e stamp of dark mottled skin in the middle
of the %hite elly1 &nd no% for the point of the story1 *ormally0 if you tickle a fro! on its ack %ith
a ristle0 the fro! %ill %ipe the place %ith a foot0 as if deterrin! an irritatin! fly1 But %hen Sperry
tickled his experimental fro! on the %hite (posta!e stamp( on its ack0 it %iped its ellyK &nd %hen
Sperry tickled it on the dark posta!e stamp on its elly0 the fro! %iped its ack1
What happens in normal emryonic development0 accordin! to Sperry(s interpretation0 is
that axons 7lon! (%ires(0 each one a narro%0 tuular extension of a sin!le nerve cell8 !ro% Euestin!ly
out from the spinal cord0 sniffin! like a do! for elly skin1 Other axons !ro% out from the spinal
cord0 sniffin! out ack skin1 &nd normally this !ives the ri!ht resultA tickles on the ack feel as
thou!h they are on the ack0 %hile tickles on the elly feel as thou!h they are on the elly1 But in
Sperry(s experimental fro!0 some of the nerve cells sniffin! out elly skin found the posta!e stamp
of elly skin !rafted on the ack0 presumaly ecause it smelled ri!ht1 &nd vice versa1 People %ho
elieve in some sort of tab"la rasa theory : %herey %e are all orn %ith a lank sheet for a mind0
and fill it in y experience : must e surprised at Sperry(s result1 They %ould expect that fro!s
%ould learn from experience to feel their %ay around their o%n skin0 associatin! the ri!ht
sensations %ith the ri!ht places on the skin1 #nstead0 it seems that each nerve cell in the spinal cord
is laelled0 say0 a elly nerve cell or a ack nerve cell0 even efore it makes contact %ith the
appropriate skin1 #t %ill later find its desi!nated tar!et pixel of skin0 %herever it may e1 #f a fly
%ere to cra%l up the len!th of its ack0 Sperry(s fro! %ould presumaly experience the illusion that
the fly suddenly leaped from ack to elly0 cra%led a little further0 then instantaneously leaped to
the ack a!ain1
Experiments like this led Sperry to formulate his (chemo:affinity( hypothesis0 accordin! to
%hich the nervous system %ires itself up not y follo%in! an overall lueprint ut y each
individual axon seekin! out end or!ans %ith %hich it has a particular chemical affinity1 Once a!ain0
%e have small0 local units follo%in! local rules1 Cells in !eneral ristle %ith (laels(0 chemical
ad!es that enale them to find their (partners(1 &nd %e can !o ack to the ori!ami analo!y to find
another place %here the laellin! principle comes in useful1 )uman paper ori!ami doesn(t use !lue0
ut it could1 &nd the ori!ami of the emryo0 %herey animal odies put themselves to!ether0 does
indeed use somethin! eEuivalent to !lue1 Glue s0 rather0 ecause there are lots of them0 and this is
%here laellin! comes triumphantly into its o%n1 Cells have a complicated repertoire of (adhesion
molecules( on their surfaces0 %herey they stick to other cells1 This cellular !luein! plays an
important role in emryonic development in all parts of the ody1 There is a si!nificant difference
from the !lues that %e are familiar %ith0 ho%ever1 ,or us0 !lue is !lue is !lue1 Some !lues are
stron!er than others0 and some !lues set faster than others0 and some !lues are more suitale for
%ood0 say0 %hile others %ork etter for metals or plastics1 But that(s aout it for variety amon!
Cell adhesion molecules are much more in!enious than that1 "ore fussy0 you could say1
$nlike our artificial !lues0 %hich %ill stick to most surfaces0 cell adhesion molecules ind only to
particular other cell adhesion molecules of exactly the ri!ht kind1 One class of adhesion molecules
in verterates0 the cadherins0 come in aout ei!hty currently kno%n flavours1 With some exceptions0
each of these ei!hty or so cadherins %ill ind only to its o%n kind1 ,or!et !lue for a minuteA a etter
analo!y mi!ht e the children(s party !ame %here each child is assi!ned an animal0 and they all
have to mill aout the room makin! noises like their o%n allotted animals1 Each child kno%s that
only one other child has een assi!ned the same animal as herself0 and she has to find her partner y
listenin! throu!h the cacophony of farmyard imitations1 Cadherins %ork like that1 Perhaps0 like me0
you can dimly ima!ine ho% the Hudicious dopin! of cell surfaces %ith particular cadherins at
strate!ic spots mi!ht refine and complicate the self:assemly principles of emryo ori!ami1 *ote0
once a!ain0 that this doesn(t imply any kind of overall plan0 ut rather a piecemeal collection of
local rules1
)avin! seen ho% %hole sheets of cells play the ori!ami !ame in shapin! the emryo0 let(s
no% dive inside a sin!le cell0 %here %e(ll find the same principle of self:foldin! and self:crumplin!0
ut on a much smaller scale0 the scale of the sin!le protein molecule1 Proteins are immensely
important0 for reasons that # must take time to explain0 e!innin! %ith a teasin! speculation to
celerate the uniEue importance of proteins1 # love speculatin! on ho% %eirdly different %e should
expect life to e else%here in the universe0 ut one or t%o thin!s # suspect are universal0 %herever
life mi!ht e found1 &ll life %ill turn out to have evolved y a process related to 'ar%inian natural
selection of !enes1 &nd it %ill rely heavily on proteins : or molecules %hich0 like proteins0 are
capale of foldin! themselves up into a hu!e variety of shapes1 Protein molecules are virtuosos of
the auto:ori!amic arts0 on a scale much smaller than that of the sheets of cells %e have so far dealt
%ith1 Protein molecules are daFFlin! sho%cases of %hat can e achieved %hen local rules are
oeyed on a local scale1
Proteins are chains of smaller molecules called amino acids0 and these chains0 like the sheets
of cells %e have een considerin!0 also fold themselves0 in hi!hly determined %ays ut on a much
smaller scale1 #n naturally occurrin! proteins 7this is one fact that %ill presumaly e different on
alien %orlds8 there are only t%enty kinds of amino acid0 and all proteins are chains strun! to!ether
from Hust this repertoire of t%enty0 dra%n from a much lar!er set of possile amino acids1 *o% for
the auto:ori!ami1 Protein molecules0 simply follo%in! the la%s of chemistry and thermodynamics0
spontaneously and automatically t%ist themselves into precisely shaped three:dimensional
confi!urations : # almost said (knots( ut0 unlike ha!fish 7if # mi!ht impart a !ratuitously
inconseEuential ut en!a!in! fact80 proteins don(t literally tie themselves in knots1 The three:
dimensional structure into %hich a protein chain folds and t%ists itself is the (tertiary structure( that
%e riefly met %hen considerin! the self:assemly of viruses1 &ny !iven seEuence of amino acids
dictates a particular foldin! pattern1 The amino:acid seEuence0 %hich itself is determined y the
seEuence of letters in the !enetic code0 determines the shape of the tertiary pattern1 L The shape of
the tertiary structure0 in turn0 has hu!ely important chemical conseEuences1
The auto:ori!ami y %hich protein chains fold and coil themselves is ruled y the la%s of
chemical attraction0 and the la%s determinin! the an!les at %hich atoms ind to one another1
#ma!ine a necklace of curiously shaped ma!nets1 The necklace %ould not han! in a !raceful
catenary around a !raceful neck1 #t %ould assume some other shape0 ecomin! tan!led up as the
ma!nets latched on to each other and slotted into each other(s nooks and crannies at various points
alon! the len!th of the chain1 $nlike the case of the protein chain0 the exact shape of the tan!le
%ould not e predictale0 ecause any ma!net %ill attract any other1 But it does su!!est ho% chains
of amino acids can spontaneously form a complicated knot:like structure0 %hich may not look like a
chain or a necklace1
The details of ho% the la%s of chemistry determine the tertiary structure of a protein are not
yet fully understoodA chemists can(t yet deduce0 in all cases0 ho% a !iven seEuence of amino acids
%ill coil up1 *evertheless0 there is !ood evidence that the tertiary structure is in principle deducile
from the seEuence of amino acids1 There(s nothin! mysterious aout the phrase (in principle(1
*oody can predict ho% a die %ill fall0 ut %e all elieve it is %holly determined y precise details
of ho% it is thro%n0 plus some additional facts aout %ind resistance and so on1 #t is a demonstrated
fact that a particular seEuence of amino acids al%ays does coil up into a particular shape0 or one of a
discrete set of alternative shapes 7see the lon! footnote opposite81 &nd : the important point for
evolution : the seEuence of amino acids is itself fully determined0 throu!h implementation of the
rules of the !enetic code0 y the seEuence of 7triplets of8 (letters( in a !ene1 Even thou!h it is not
easy for human chemists to predict %hat chan!e in protein shape %ill result from a particular
!enetic mutation0 it is still a fact that once a mutation has occurred0 the resultin! chan!e of protein
shape %ill e in principle predictale1 The same mutant !ene %ill relialy produce the same altered
protein shape 7or discrete menu of alternative shapes81 &nd that is all that matters for natural
selection1 *atural selection doesn(t need to understand %hy a !enetic chan!e has a certain
conseEuence1 #t is sufficient that it does1 #f that conseEuence affects survival0 the chan!ed !ene itself
%ill stand or fall in the competition to dominate the !ene pool0 %hether or not %e understand the
exact route y %hich the !ene affects the protein1
Given that protein shape is immensely versatile0 and !iven that it is determined y !enes0
%hy is it so supremely importantB Partly ecause some proteins have a direct structural role to play
in the ody1 ,irous proteins0 such as colla!en0 Hoin to!ether in stout ropes0 %hich %e call li!aments
and tendons1 But most proteins are not firous1 #nstead0 they fold themselves up into their o%n
characteristic !loular shape0 complete %ith sutle dents0 and this shape determines the protein(s
characteristic role as an en2yme0 %hich is a catalyst1
& catalyst is a chemical sustance that speeds up0 y as much as a illion or even a trillion
times0 a chemical reaction et%een other sustances0 %hile the catalyst itself emer!es from the
process unscathed and free to catalyse a!ain1 EnFymes0 %hich are protein catalysts0 are champions
amon! catalysts ecause of their specificityA they are very fussy aout precisely %hich chemical
reactions they speed up1 Or perhaps %e could sayA chemical reactions in livin! cells are very fussy
aout %hich enFymes speed them up1 "any reactions in cell chemistry are so slo% that0 %ithout the
ri!ht enFyme0 for practical purposes they don(t occur at all1 But %ith the ri!ht enFyme they happen
very fast0 and can churn out products in ulk1
)ere(s ho% # like to put it1 & chemistry la has hundreds of ottles and Hars on its shelves0
each containin! a different pure sustanceA compounds and elements0 solutions and po%ders1 &
chemist %ishin! to perform a certain chemical reaction selects t%o or three ottles0 takes a sample
from each0 mixes them in a test tue or a flask0 perhaps applies heat0 and the reaction takes place1
Other chemical reactions that could take place in the la don(t0 ecause the !lass %alls of the ottles
and Hars prevent the in!redients meetin!1 #f you %ant a different chemical reaction0 you mix
different in!redients in a different flask1 Every%here there are !lass arriers keepin! the pure
sustances separate from one another in ottles or Hars0 and keepin! the reactin! cominations
separate from one another in test tues or flasks or eakers1
The livin! cell0 too0 is a !reat chemistry la0 and it has a similarly lar!e store of chemicals1
But they aren(t kept in separate ottles and Hars on shelves1 They are all mixed up to!ether1 #t is as
thou!h a vandal0 a chemical lord of misrule0 entered the la0 seiFed all the ottles on all the shelves0
and tipped them %ith anarchistic aandon into one !reat cauldron1 Terrile thin! to doB Well0 it
%ould e if they all reacted to!ether0 in all possile cominations1 But they don(t1 Or if they do0 the
rate at %hich they react to!ether is so slo% that they mi!ht as %ell not e reactin! at all1 Except :
and this is the %hole point : if an enFyme is present1 There is no need for !lass ottles and Hars to
keep the sustances apart ecause0 to all intents and purposes0 they are not !oin! to react to!ether
any%ay : "nless the ri!ht enFyme is present1 The eEuivalent of keepin! the chemicals in stoppered
ottles until you %ant to mix a particular pair0 say & and B0 is to mix all the hundreds of sustances
up in a !reat %itch(s re%0 ut supply only the ri!ht enFyme to catalyse the reaction et%een & and
B and no other comination1 &ctually0 the metaphor of the anarchically inclined ottle:tipper !oes
too far1 Cells do contain an infrastructure of memranes et%een %hich0 and %ithin %hich0
chemical reactions !o on1 To some extent0 these memranes play the role of !lass partitions
et%een test tues and flasks1
The point of this section of the chapter is that (the ri!ht enFyme( achieves its (ri!htness(
lar!ely throu!h its physical shape 7and that(s important0 ecause the physical shape is determined y
!enes0 and it is !enes %hose variations are ultimately favoured or disfavoured y natural selection81
"olecules aplenty are driftin! and t%istin! and spinnin! throu!h the soup that athes the interior of
a cell1 & molecule of sustance & mi!ht e happy to react %ith a molecule of sustance B0 ut only
if they happen to collide %hen facin! in exactly the ri!ht direction0 relative to each other1 Crucially0
that seldom happens : "nless the ri!ht enFyme intervenes1 The enFyme(s precise shape0 the shape
into %hich it folded itself like a ma!netic necklace0 leaves it pitted %ith cavities and dents0 each one
of %hich itself has a precise shape1 Each enFyme has a so:called (active site(0 %hich is usually a
particular dent or pocket0 %hose shape and chemical properties confer upon the enFyme its
specificity1 The %ord (dent( doesn(t adeEuately convey the specificity0 the precision0 of this
mechanism1 Perhaps a etter comparison is %ith an electric socket1 #n %hat my friend the Foolo!ist
.ohn Dres calls (the !reat plu! conspiracy(0 different countries around the %orld have irritatin!ly
adopted different aritrary conventions for plu!s and sockets1 British plu!s %on(t fit &merican
sockets0 or ,rench sockets0 and so on1 The active sites on the surface of protein molecules are
sockets into %hich only certain molecules %ill fit1 But %hereas the !reat plu! conspiracy runs to
only half a doFen separate shapes around the %orld 7Euite enou!h to constitute a continual
annoyance to the traveller80 the different kinds of sockets sported y enFymes are far more
Think of a particular enFyme0 %hich catalyses the chemical comination of t%o molecules0
P and P0 to make the compound PP1 One half of the active site (socket( is Hust ri!ht for a molecule
of type P to nestle into0 like a Hi!sa% piece1 The other half of the same socket is eEually precisely
shaped for a P molecule to slot in : facin! exactly the ri!ht %ay to comine chemically %ith the P
molecule that is already there1 Sharin! a dent0 firmly held at Hust the ri!ht an!le to each other y the
matchmakin! enFyme molecule0 P and P unite1 The ne% compound0 PP0 no% reaks a%ay into the
soup0 leavin! the active dent in the enFyme molecule free to rin! to!ether another P and another P1
& cell may e filled %ith s%arms of identical enFyme molecules0 all %orkin! a%ay like roots in a
car factory0 churnin! out PP in the cellular eEuivalent of industrial Euantities1 Put a different
enFyme into the same cell0 and it %ill churn out a different product0 perhaps PR0 or PS or 6I1 The
end product is different0 even thou!h the availale ra% materials are the same1 Other types of
enFymes are concerned not %ith constructin! ne% compounds0 ut %ith reakin! do%n old ones1
Some of these enFymes are involved in di!estin! food0 and they are exploited0 too0 in (iolo!ical(
%ashin! po%ders1 But0 since this chapter is aout the construction of emryos0 %e are here mostly
concerned %ith constructive enFymes0 %hich roker the synthesis of ne% chemical compounds1 One
such process is sho%n in action on colour pa!e 231
& prolem may have occurred to you1 #t(s all very %ell to talk of Hi!sa% dents and sockets0
hi!hly specific active sites capale of speedin! up a particular chemical reaction a trillionfold1 But
doesn(t it sound too !ood to e trueB )o% do enFyme molecules of exactly the ri!ht shape evolve
from less perfect e!innin!sB What is the proaility that a socket0 shaped at random0 %ill have Hust
the ri!ht shape0 and Hust the ri!ht chemical properties0 to arran!e a marria!e et%een t%o molecules0
P and P0 finessin! their encounter at exactly the ri!ht an!leB *ot very !reat if you think (finished
Hi!sa%( : or0 indeed0 if you think (!reat plu! conspiracy(1 #nstead0 you have to think (smooth !radient
of improvement(1 &s so often %hen %e are faced %ith the riddle of ho% complex and improale
thin!s can arise in evolution0 it is a fallacy to assume that the final perfection that %e see today is
the %ay it al%ays %as1 ,ully fashioned0 hi!hly evolved enFyme molecules achieve trillionfold
speedups of the reactions they catalyse0 and they do so y ein! eautifully crafted to exactly the
ri!ht shape1 But you don(t need a trillionfold speedup in order to e favoured y natural selection1 &
millionfold %ill do nicelyK So %ill a thousandfold1 &nd even tenfold or t%ofold %ould e enou!h
for natural selection to !et an adeEuate !rip1 There is a smooth !radient of improvement in an
enFyme(s performance0 all the %ay from almost no dent at all0 throu!h a crudely shaped dent0 to a
socket of exactly the ri!ht shape and chemical si!nature1 (Gradient( means that each step is a
noticeale improvement0 ho%ever sli!ht0 over the one efore1 &nd (noticeale( for natural selection
can mean an improvement smaller than the minimum that %ould e reEuired for us to notice it1
So0 you see the %ay it %orks1 Ele!antK & cell is a versatile chemical factory0 capale of
spe%in! out massive Euantities of a %ide variety of different sustances0 the choice ein! made y
%hich enFyme is present1 &nd ho% is that choice madeB By %hich !ene is t"rned on1 .ust as the cell
is a vat filled %ith lots of chemicals0 only a minority of %hich react %ith each other0 so every cell
nucleus contains the entire !enome0 ut %ith only a minority of !enes turned on1 When a !ene is
turned on in0 say0 a cell of the pancreas0 its seEuence of code letters directly determines the
seEuence of amino acids in a proteinJ and the seEuence of amino acids determines 7rememer the
ima!e of the ma!netic necklaceB8 the shape into %hich the protein folds itselfJ and the shape into
%hich the protein folds itself determines the precisely shaped sockets that marry up sustances
driftin! around in the cell1 Every cell0 %ith very fe% exceptions such as red lood corpuscles0 %hich
lack a nucleus0 contains the !enes for makin! all the enFymes1 But in any one cell0 only a fe% !enes
%ill e turned on at any one time1 #n0 say0 thyroid cells0 the !enes that make the ri!ht enFymes for
catalysin! the manufacture of thyroid hormone are turned on1 &nd correspondin!ly for all the
different kinds of cells1 ,inally0 the chemical reactions that !o on in a cell determine the %ay that
cell is shaped and the %ay it ehaves0 and the %ay it participates in ori!ami:style interactions %ith
other cells1 So the %hole course of emryonic development is controlled0 via an intricate seEuence
of events0 y !enes1 #t is !enes %hich determine seEuences of amino acids0 %hich determine tertiary
structures of proteins0 %hich determine the socket:like shapes of active sites0 %hich determine cell
chemistry0 %hich determine (starlin!:like( cell ehaviour in emryonic development1 So0 differences
in !enes can0 at the ori!inatin! end of the complex chain of events0 cause differences in the %ay
emryos develop0 and hence differences in the form and ehaviour of adults1 The survival and
reproductive success of those adults then feeds ack on the survival in the !ene pool of the !enes
that made the difference et%een success and failure1 &nd that is natural selection1
Cell(lar $a"ily tree o$#aenorha$ditis elegans
Emryolo!y seems complicated : is complicated : ut it is easy to !rasp the important point0
%hich is that %e are dealin! %ith local self:assemly processes all the %ay1 #t(s a separate Euestion0
!iven that 7almost8 all the cells contain all the !enes0 ho% it is decided %hich !enes are turned on in
each different kind of cell1 # must riefly deal %ith that no%1
T)E* WOR"S S)&-- TR6
Whether or not a !iven !ene is turned on in a !iven cell at a !iven time is determined0 often
via a cascade of other !enes called s%itch !enes or controller !enes0 y the chemical environment
of the cell1 Thyroid cells are Euite different from muscle cells0 and so on0 even thou!h their !enes
are the same1 That(s all very %ell0 you may say0 once the development of the emryo is under %ay0
and the different kinds of tissues such as thyroid and muscle already exist1 But every emryo starts
out as a sin!le cell1 Thyroid cells and muscle cells0 liver cells and one cells0 pancreas cells and skin
cells0 all are descended from a sin!le fertiliFed e!! cell0 via a ranchin! family tree1 This is a
cellular family tree !oin! ack no further than the moment of conception0 nothin! to do %ith the
evolutionary tree !oin! ack millions of years0 %hich keeps croppin! up in other chapters1 -et me
sho% you0 for example0 the complete family tree of all ==; cells of a ne%ly hatched larva of the
nematode %orm0 #aenorhabditis ele$ans 7elo%A please pay close attention to every detail of this
dia!ram81 By the %ay0 # don(t kno% %hat this tiny %orm did to earn its species name of ele$ans0 ut
# can think of a !ood reason %hy it mi!ht have deserved it retrospectively1 # kno% that not all my
readers like my di!ressions0 ut the research that has een done on #aenorhabditis ele$ans is such a
rin!in! triumph of science that you aren(t !oin! to stop me1
#aenorhabditis ele$ans %as chosen in the 29<5s as an ideal experimental animal y the
formidaly rilliant South &frican iolo!ist Sydney Brenner1 )e had recently completed his %ork0
%ith ,rancis Crick and others at Camrid!e0 on crackin! the !enetic code0 and %as lookin! around
for a ne% i! prolem to solve1 )is inspired choice0 and his o%n pioneerin! research on its !enetics
and neuro:anatomy0 has led to a %orld%ide community of #aenorhabditis researchers that has
!ro%n into the thousands1 #t is only a it of an exa!!eration to say that %e no% kno% eerythin$
aout #aenorhabditis ele$ansK We kno% its entire !enome1 We kno% exactly %here every one of
its ==; cells 7in the larvaJ 9=9 in the adult hermaphroditic form0 not countin! reproductive cells8 is
in the ody0 and %e kno% the exact (family history( of every one of those cells0 throu!h emryonic
development1 We kno% of a lar!e numer of mutant !enes0 %hich produce anormal %orms0 and
%e kno% exactly %here the mutation acts in the ody and the exact cellular history of ho% the
anormality develops1 This little animal is kno%n from start to finish0 kno%n inside out0 kno%n
from head to tail and all stations in et%een0 kno%n throu!h and throu!h 7(O fraHous dayK(81
Brenner %as elatedly reco!niFed %ith the *oel PriFe for Physiolo!y in 35530 and a related
species %as named in his honour0 #aenorhabditis brenneri1 )is re!ular column in the Hournal
#"rrent Biolo$y0 under the yline ($ncle Syd(0 is a model of intelli!ent and irreverent scientific %it
: as ele!ant as the %orld%ide research effort on #4 ele$ans that he inspired1 But # do %ish molecular
iolo!ists %ould talk to some Foolo!ists 7like Brenner himself8 and learn not to refer to
#aenorhabditis as (the( nematode0 or even (the( %orm0 as thou!h there %ere no others1
Of course you can(t read the names of the cell types at the ottom of the dia!ram 7it %ould
take seven pa!es to print the %hole thin! out le!ily80 ut they say thin!s like (pharynx(0 (intestinal
muscle(0 (ody muscle(0 (sphincter muscle(0 (rin! !an!lion(0 (lumar !an!lion(1 The cells of all these
types are literally cousins of one anotherA cousins y virtue of their ancestry %ithin the lifetime of
the individual %orm1 ,or example0 # am lookin! at a particular ody muscle cell called "Spappppa0
%hich is a silin! of another ody muscle cell0 first cousin of t%o more ody muscle cells0 first
cousin once removed of t%o more ody muscle cells0 second cousin of six pharynx cells0 third
cousin of seventeen pharynx cells 1 1 1 and so on1 #sn(t it amaFin! that %e can actually use %ords like
(second cousin once removed(0 %ith the utmost precision and certainty0 to refer to named and
repeataly identifiale cells in an animal(s odyB The numer of cell (!enerations( that separates the
tissues from the ori!inal e!! is not that !reat1 &fter all0 there are only ==; cells in the ody0 and you
can theoretically make 2053> 73 to the po%er 258 in ten !enerations of cell splittin!1 The numers of
cell !enerations for human cells %ould e much lar!er1 *evertheless0 you could in theory make a
similar family tree for every one of your trillion:odd cells 7as opposed to the ==; cells of a #4
ele$ans female larva80 tracin! each one(s descent ack to the one fertiliFed e!! cell1 #n mammals0
ho%ever0 it is not possile to identify particular0 repeataly named cells1 #n us0 it is more a case of
statistical populations of cells0 %hose details are different in different people1
# hope my euphoric di!ression on the ele!ance of #aenorhabditis research has not distracted
us too far from the point # %as makin! aout ho% cell types chan!e in their shape and character as
they ranch a%ay from one another in the emryonic family tree1 &t the ranchin! point et%een a
clone that is destined to ecome pharynx cells0 and a (cousin( clone that is destined to ecome rin!
!an!lion cells0 there has to e somethin! to distin!uish them0 other%ise ho% %ould they kno% to
turn on different !enesB The ans%er is that0 %hen the most recent common ancestor of the t%o
clones divided0 the t%o halves of the cell efore division %ere different1 So0 %hen the cell divided0
the t%o dau!hter cells0 thou!h identical in their !enes 7every dau!hter cell receives a full
complement of !enes80 %ere not identical in the surroundin! chemicals1 &nd this meant that the
same !enes %ere not turned on : %hich chan!ed the fate of their descendants1 The same principle
applies ri!ht throu!h emryolo!y0 includin! its very start1 The key to differentiation0 in all animals0
is asymmetric cell division1 L
Sir .ohn Sulston and his collea!ues traced each of the cells in the ody of the %orm ack to
one and only one of six founder cells : %e mi!ht even call them (matriarch( cells : called &B0 "S0 E0
'0 C and P>1
#n namin! the cells0 they used a neat notation that summariFed the history of each
one1 Every cell(s name e!ins %ith the name of one of those six founder cells0 the one from %hich it
is descended1 Thereafter0 its name is a strin! of letters0 the initial letters of the direction of cell
division that !ave rise to itA anterior0 posterior0 dorsal0 ventral0 left0 ri!ht1 ,or example0 Ca and Cp
are the t%o dau!hters of matriarch C0 the anterior and posterior dau!hter respectively1 *otice that
every cell has no more than t%o dau!hters 7of %hich one may die81 # am no% lookin! at a particular
ody muscle cell0 %hose name0 Cappppv0 succinctly discloses its historyA C had an anterior
dau!hter0 %hich had a posterior dau!hter0 %hich had a posterior dau!hter0 %hich had a posterior
dau!hter0 %hich had a posterior dau!hter0 %hich had a ventral dau!hter0 %hich is the ody muscle
cell in Euestion1 Every cell in the ody is denoted y a comparale strin! of letters headed y one of
the six founder cells1 &Bprpapppap0 to take another example0 is a nerve cell that sits in the ventral
nerve cord runnin! alon! the len!th of the %orm1 *eedless to say0 it is not necessary to take in the
details1 The eautiful point is that every sin!le cell in the ody has such a name0 %hich totally
descries its history durin! emryolo!y1 Every one of the ten cell divisions that !ave rise to
&Bprpapppap0 and every other cell0 %as an asymmetric division %ith the potential for different
!enes to e s%itched on in each of the t%o dau!hter cells1 &nd in all animals that is the principle y
%hich tissues differentiate0 even thou!h all their cells contain the same !enes1 "ost animals0 of
course0 have far more cells than #aenorhabditis' ==;0 and their emryonic development is in most
cases less ri!idly determined1 #n particular0 as Sir .ohn Sulston kindly reminds me0 and as # have
already riefly mentioned0 in a mammal the (family trees( of our cells are different for every
individual0 %hereas in #aenorhabditis they are almost identical 7except in mutant individuals81
*evertheless0 the principle remains the same1 #n any animal0 cells differ from each other in different
parts of the ody0 even thou!h they are !enetically identical0 ecause of their history of asymmetric
cell division durin! the short course of emryonic development1
-et us hear the conclusion of the %hole matter1 There is no overall plan of development0 no
lueprint0 no architect(s plan0 no architect1 The development of the emryo0 and ultimately of the
adult0 is achieved y local rules implemented y cells0 interactin! %ith other cells on a local asis1
What !oes on inside cells0 similarly0 is !overned y local rules that apply to molecules0 especially
protein molecules0 %ithin the cells and in the cell memranes0 interactin! %ith other such
molecules1 &!ain0 the rules are all local0 local0 local1 *oody0 readin! the seEuence of letters in the
'*& of a fertiliFed e!!0 could predict the shape of the animal it is !oin! to !ro% into1 The only
%ay to discover that is to !ro% the e!!0 in the natural %ay0 and see %hat it turns into1 *o electronic
computer could %ork it out0 unless it %as pro!rammed to simulate the natural iolo!ical process
itself0 in %hich case you mi!ht as %ell dispense %ith the electronic version and use the developin!
emryo as its o%n computer1 This %ay of !eneratin! lar!e and complex structures purely y the
execution of local rules is deeply distinct from the lueprint %ay of doin! thin!s1 #f the '*& %ere
some kind of lineariFed lueprint0 it %ould e a relatively trivial exercise to pro!ram a computer to
read the letters and dra% the animal1 But it %ould not e at all easy : indeed0 it mi!ht e impossile
: for the animal to have evolved in the first place1
&nd no%0 so that this chapter on emryos should not end up as a mere di!ression in a ook
on evolution0 # must return to the sincere dilemma of )aldane(s Euestioner1 Given that !enes control
processes of emryonic development rather than adult shapeJ !iven that natural selection : like God
: doesn(t uild tiny %in!s0 ut emryolo!y doesJ ho% does natural selection !o to %ork on animals
to shape their odies and their ehaviourB )o% does natural selection !o to %ork on emryos0 in
other %ords0 to reHi! them so they ecome ever more proficient at uildin! successful odies0 %ith
%in!s0 or fins0 leaves or armour platin!0 stin!s or tentacles or %hatever it takes to surviveB
*atural selection is the differential survival of successful !enes rather than alternative0 less
successful !enes in !ene pools1 *atural selection doesn(t choose !enes directly1 #nstead it chooses
their proxies0 individual odiesJ and those individuals are chosen : oviously and automatically and
%ithout delierative intervention : y %hether they survive to reproduce copies of the very same
!enes1 & !ene(s survival is intimately ound up %ith the survival of the odies that it helps to uild0
ecause it rides inside those odies0 and dies %ith them1 &ny !iven !ene can expect to find itself0 in
the form of copies of itself0 ridin! inside a lar!e numer of odies0 oth simultaneously in a
population of contemporaries0 and successively as !eneration !ives %ay to !eneration1 Statistically0
therefore0 a !ene that tends0 on avera!e0 to have a !ood effect on the survival prospects of the
odies in %hich it finds itself %ill tend to increase in freEuency in the !ene pool1 So0 on avera!e0 the
!enes that %e encounter in a !ene pool %ill tend to e those !enes that are !ood at uildin! odies1
This chapter has een aout the procedures y %hich !enes uild odies1
)aldane(s interlocutor found it implausile that natural selection could put to!ether in0 say0 a
illion years0 a !enetic recipe for uildin! her1 # find it plausile0 althou!h of course neither # nor
anyody else can tell you the details of ho% it happened1 The reason it is plausile is precisely that
it is all done y local rules1 #n any one act of natural selection0 the mutation that is selected has had :
in lots of cells and in lots of individuals in parallel : a very simple effect on the shape into %hich a
protein chain spontaneously coils up1 This0 in turn0 throu!h catalytic action0 speeds up0 say0 a
particular chemical reaction in all the cells in %hich the !ene is turned on1 This chan!es0 perhaps0
the rate of !ro%th of the emryonic primordium of the Ha%1 &nd this has conseEuential effects on
the shape of the %hole face0 perhaps shortenin! the muFFle and !ivin! a more human and less (ape:
like( profile1 *o%0 the natural selection pressures that favour or disfavour the !ene can e as
complicated as you like1 They mi!ht involve sexual selection0 perhaps aesthetic choice of a hi!h
order y %ould:e sexual partners1 Or the chan!e in Ha% shape mi!ht have a sutle effect on the
animal(s aility to crack nuts0 or its aility to fi!ht rivals1 Some hu!ely elaorate comination of
selection pressures0 conflictin! and compromisin! %ith one another in e%ilderin! complexity0 can
ear upon the statistical success of this particular !ene0 as it propa!ates itself throu!h the !ene pool1
But the !ene kno%s nothin! of this1 &ll it is doin!0 %ithin different odies and in successive
!enerations0 is reHi!!in! a carefully sculpted dent in a protein molecule1 The rest of the story
follo%s automatically0 in ranchin! cascades of local conseEuences0 from %hich0 eventually0 a
%hole ody emer!es1
Even more complicated than the selection pressures in the ecolo!ical0 sexual and social
environments of the animals is the phantasma!oric net%ork of influences that !o on %ithin and
amon! the developin! cellsA influences of !enes on proteins0 !enes on !enes0 proteins on the
expression of !enes0 proteins on proteinsJ memranes0 chemical !radients0 physical and chemical
!uide rails in emryos0 hormones and other mediators of action at a distance0 laelled cells seekin!
others %ith identical or complementary laels1 *oody understands the %hole picture0 and noody
needs to understand it in order to accept the exEuisite plausiility of natural selection1 *atural
selection favours the survival in the !ene pool of the !enetic mutations responsile for makin!
crucial chan!es in emryos1 The %hole picture emer!es as a conseEuence of hundreds of thousands
of small0 local interactions0 each one comprehensile in principle 7althou!h it may e too hard or
too time:consumin! to unravel in practice8 to anyone %ith sufficient patience to examine it1 The
%hole may e afflin! and mysterious in practice0 ut there is no mystery in principle0 either in
emryolo!y itself0 or in the evolutionary history y %hich the controllin! !enes came to
prominence in the !ene pool1 The complications accumulated !radually over evolutionary timeA
each step %as only a tiny it different from the one efore0 and each step %as accomplished y a
small0 sutle chan!e in an existin! local rule1 When you have a sufficient numer of small entities :
cells0 protein molecules0 memranes : each at its o%n level oeyin! local rules and influencin!
others : then the eventual conseEuence is dramatic1 #f !enes survive or fail to survive as a
conseEuence of their influence on such local entities and their ehaviour0 natural selection of
successful !enes : and the emer!ence of their successful products : %ill inevitaly follo%1 )aldane(s
Euestioner %as %ron!1 #t is not in principle difficult to make somethin! like her1
&nd0 as )aldane said0 it only takes nine months1
L # have een %arned that (&ll thin!s ri!ht and eautiful( %ill not necessarily strike my
readers as nostal!ically as it does me1 #t is an &n!lican hymn for children %ritten y "rs C1 ,1
&lexander in 2;>;0 comfortaly extollin! the eauties of nature 7and0 in one verse0 the political
status Euo8 %ith the refrain0 (The -ord God made them all(1 #t is the suHect of a splendid parody
%ritten y Eric #dle and sun! y the "onty Python teamA
&ll thin!s dull and u!ly
&ll creatures short and sEuat
&ll thin!s rude and nasty
The -ord God made the lot1
Each little snake that poisons
Each little %asp that stin!s
)e made their rutish venom
)e made their horrid %in!s1
&ll thin!s sick and cancerous
&ll evil !reat and small0
&ll thin!s foul and dan!erous
The -ord God made them all1
Each nasty little hornet
Each eastly little sEuid
Who made the spiky urchinB
Who made the sharksB )e didK
&ll thin!s scaed and ulcerous
&ll pox oth !reat and small
Putrid0 foul and !an!renous
The -ord God made them all1
L *ote for professionals at the interface et%een iolo!ists and computer scientistsA Charles
Simonyi0 %ho speaks %ith the authority of a distin!uished soft%are desi!ner0 put it as follo%s0 after
readin! an early draft of this chapterA (1 1 1 the recipe 7of the eye0 rain0 lood0 etc18 is much much
simpler than the lueprint for the same 7in terms of its or ase:pairs8 so evolution %ould e
literally impossile 7in less than 25T255 years8 especially ecause small variations in the lueprint
%ould e unlikely to have any positive effect0 %hereas a variation in the recipe %ould1( &lludin! to
my o%n (computer iomorphs( and (arthromorphs( 7see Chapter 380 'r Simonyi !oes onA (The
artificial creatures that you Mpro!rammed for The Blind Watchmaker and #limbin$ %o"nt
&mprobableN are all descried y recipes0 not y lueprint : a lueprint %ould e Hust a Humle of
vectors of lack lines : can you ima!ine tryin! to play evolution on them y varyin! the endpoints
of the lack lines one at a time or even t%o at a timeB( &s you(d expect from one descried y Bill
Gates0 no less0 as (one of the !reat pro!rammers of all time(0 this is exactly ri!ht for the computer
iomorphs0 and it is surely ri!ht for livin! thin!s too1
L There is a risk that (epi!enesis( %ill e confused %ith (epi!enetics(0 a modish uFF:%ord
no% enHoyin! its fifteen minutes of fame in the iolo!ical community1 Whatever (epi!enetics( mi!ht
mean 7and its enthusiasts cannot seem to a!ree even %ith themselves0 let alone %ith each other80 all
# intend to say aout it here is that it is not the same thin! as epi!enesis1
L "y medieval historian collea!ue 'r Christopher Tyerman confirms that this %as indeed a
myth that %as invented in +ictorian times for idealistic reasons0 ut that there %as never a scintilla
of truth in it1
L #nva!inateA (fold in%ards to form a hollo%(0 (turn or doule ack %ithin itself( 7 Shorter
!xford En$lish Dictionary81
L The craFe died out0 ut # reintroduced it to the same school in the 29=5s0 %hereupon it
spread Hust like a second epidemic of the same disease1
L # am sorry # am at a loss to explain %hy the notochord !ets an (h(0 like a musical or
mathematical chord0 %hile the spinal cord doesn(t0 like a it of strin!1 # have al%ays found it
mysterious0 and have even %ondered %hether it mi!ht represent some lon!:for!otten ut fossiliFed
mistake1 &dmittedly0 the !xford En$lish Dictionary lists (chord( as an alternative spellin! for the
strin! kind of cord0 ut the difference does seem Eueer !iven that the spinal cord and the notochord
run the len!th of the emryonic ody0 one aove the other1
L &nd that(s a fascinatin! story in its o%n ri!ht0 y the %ay1 #t has !ripped my ima!ination
ever since the !reat Camrid!e physiolo!ist .oseph *eedham 7a polymath %ho ecame even etter
kno%n as the leadin! expert on the history of Chinese science8 came to my school to demonstrate it0
at the invitation of his nephe% %ho happened to e our student teacher at the timeA a oon of
nepotism for %hich # remain !rateful1 $nder 'r *eedham(s !uidance0 %e peered at muscle fires
do%n our microscopes and %atched them shorten0 as if y ma!ic0 %hen %e !ave them a drop of
&TP0 adenosine triphosphate0 the universal ener!y currency of the ody1
L This statement needs an important reservation1 The determination of the amino:acid
seEuence y !enes is indeed asolute1 But the determination of the three:dimensional shape y the
one:dimensional amino:acid seEuence is not asolute0 and it really matters1 There are some
seEuences of amino acids that are capale of coilin! up into t%o alternative 4' shapes1 The proteins
called prions0 for example0 have t%o stale shapes1 These are discrete alternatives %ithout stale
intermediates0 in the same %ay as a li!ht s%itch is stale in the up position and in the do%n position
ut no%here in et%een1 Such (s%itch proteins( can e disastrous or they can e useful1 #n the case
of prions they are disastrous1 #n (mad co% disease(0 a useful protein in the rain 7it(s a normal
constituent of cell memranes8 happens to have an alternative form : an alternative %ay to fold
itself in auto:ori!ami1 The alternative form is normally never seen0 ut if it ever arises in one
molecule it tri!!ers nei!hourin! molecules to follo% suitA they copy it and flip to the alternative
form1 -ike a %ave of fallin! dominoes0 or like the irresponsile spreadin! of a rumour0 the
alternative prion form spreads throu!h the rain0 %ith disastrous results for the co% : or the person
in the case of CreutFfeldt:.ako disease0 or the sheep in the case of scrapie1 But sometimes
molecules %ith the aility to auto:ori!ami themselves into more than one alternative shape are
useful1 Without leavin! the metaphor of the li!ht s%itch %e find a eautiful example1 Rhodopsin0
the protein in our eyes that is responsile for our sensitivity to li!ht0 has an emedded component
called retinal 7not itself a protein8 %hich flips from its main stale confi!uration to its alternative
confi!uration %hen hit y a photon of li!ht1 #t then s%iftly reverts0 like a li!ht s%itch on a cost:
cuttin! timer0 ut mean%hile the flip has re!istered %ith the rainA (-i!ht detected at this pinpoint
location here1( .acEues "onod(s %onderful ook0 #hance and .ecessity0 is especially !ood on such
i:stale s%itch molecules1
L #n #aenorhabditis the ori!inal cell0 called I0 has a front end %hich is different from its
rear end0 and this difference %ill come to represent the eventual fore:and:aft ody axis : anterior
7front8 and posterior 7rear81 When the cell divides0 the anterior dau!hter cell0 %hich is called &B0
has more front:end sustance than the posterior dau!hter cell0 %hich is called P20 and this
difference %ill e ootstrapped to make more differences do%n the line1 &B is destined to !ive rise
to %ell over half the cells of the ody0 includin! most of the nervous system0 and # %on(t discuss it
further1 P2 has t%o children0 a!ain different from each other0 called E"S 7definin! the ventral or
elly side of the eventual %orm8 and P3 7definin! the dorsal side81 They are !randchildren of I
7rememer0 %hen # use %ords like (children( and (!randchildren(0 # am talkin! aout cells %ithin a
developin! emryo0 not individual %orms81 E"S no% has t%o children called E and "S0 %hile P3
has t%o children called C and P41 E0 "S0 C and P4 are !reat:!randchildren of I 7the other !reat:
!randchildren are descended from &B0 and # am not %ritin! them do%n0 except to say that t%o of
them0 called &Bal and &Bpl0 define the left side0 and their cousins0 &Bar and &pr0 define the ri!ht
side of the eventual %orm81 P4 has t%o children called ' and P>0 %hich are !reat:!reat:
!randchildren of I1 "S and C also have children0 ut # shan(t name them here1 P> is destined to
!ive rise to the so:called !erm line1 The !erm line consists of cells that are not involved in uildin!
the ody0 ut instead are !oin! to make the reproductive cells1 Oviously there is no need to
rememer or take note of these cell names1 The point is only that0 althou!h !enetically identical to
each other0 they differ in their chemical nature0 as a cumulatively ootstrapped conseEuence of their
history in the seEuence of cell divisions %ithin the emryo1
O Sulston0 %ho stayed at Camrid!e after Brenner left for &merica0 %as another of the
triumvirate %ho %on the *oel PriFe for the #aenorhabditis %ork1 Sulston %ent on to lead the
British end of the official )uman Genome ProHect0 the &merican end of %hich %as headed first y
.ames Watson and later y ,rancis Collins1
# "&G#*E a %orld %ithout islands1
Biolo!ists often use the %ord (island( to mean somethin! other than Hust a piece of land
surrounded y %ater1 ,rom the point of vie% of a fresh%ater fish0 a lake is an islandA an island of
haitale %ater surrounded y inhospitale land1 ,rom the point of vie% of an &lpine eetle0
incapale of flourishin! elo% a certain altitude0 each hi!h peak is an island0 %ith almost
impassale valleys et%een1 There are tiny nematode %orms 7related to the ele!ant #aenorhabditis8
%hich live inside leaves 7as many as 250555 of them in a sin!le adly infected leaf80 divin! into
them throu!h the stomata0 %hich are the microscopic holes throu!h %hich leaves take in caron
dioxide and release oxy!en1 To a leaf:d%ellin! nematode %orm such as Aphelencoides0 a sin!le
fox!love is an island1 To a louse0 a sin!le human head or crotch mi!ht e an island1 There must e
lots of animals and plants that re!ard an oasis in a desert as an island of cool0 !reen haitaility
surrounded y a hostile sea of sand1 &nd0 %hile %e are redefinin! %ords from an animal(s point of
vie%0 since an archipela!o is a chain or cluster of islands0 # suppose a fresh%ater fish mi!ht define
an archipela!o as a chain or cluster of lakes0 such as the lakes alon! the Great Rift +alley in &frica1
&n &lpine marmot mi!ht define a chain of mountain peaks separated y valleys as an archipela!o1
& leaf:minin! insect mi!ht re!ard an avenue of trees as an archipela!o1 & otfly mi!ht re!ard a
herd of cattle as a movin! archipela!o1
)avin! redefined the %ord (island( 7the saath %as made for man0 not man for the saath8
let me return to my openin!1 #ma!ine a %orld %ithout islands1
)e had ou!ht a lar!e map representin! the sea
Without the least vesti!e of landA
&nd the cre% %ere much pleased %hen they found it to e
& map they could all understand1
We %on(t !o Euite as far as the Bellman0 ut ima!ine if all the land %ere !athered to!ether
in one !reat continent in the middle of a featureless sea1 There are no islands offshore0 no lakes or
mountain ran!es on the landA nothin! to reak the monotonous s%eep of smooth uniformity1 #n this
%orld an animal can easily !o from any%here to any%here else0 limited only y sheer distance0
untrouled y inhospitale arriers1 This is not a %orld friendly to evolution1 -ife on Earth %ould
e extremely orin! if there %ere no islands0 and # %ant to e!in this chapter y explainin! %hy1
Every species is a cousin of every other1 &ny t%o species are descended from an ancestral
species0 %hich split in t%o1 ,or example0 the common ancestor of people and ud!eri!ars lived
aout 425 million years a!o1 The ancestral species split in t%o0 and the t%o strands %ent their
separate %ays for the rest of time1 # chose human and ud!ie to make it vivid0 ut that same
ancestral species is shared y all mammals on one side of that early divide0 and all reptiles
7Foolo!ically speakin!0 irds are reptiles0 as %e sa% in Chapter <8 on the other side1 #n the unlikely
event that a fossil of this ancestral species %as ever found0 it %ould need a name1 -et(s call it
Protamnio darwinii1 We don(t kno% any details aout it0 and the details don(t matter at all for the
ar!ument0 ut %e %on(t !o far %ron! if %e ima!ine it as a spra%lin! liFard:like creature0 scurryin!
aout catchin! insects1 *o%0 here(s the point1 When Protamnio darwinii split into t%o su:
populations they %ould have looked Hust the same as each other0 and could have happily interred
%ith each otherJ ut one lot %ere destined to !ive rise to the mammals0 and the other lot %ere
destined to !ive rise to the irds 7and dinosaurs and snakes and crocodiles81 These t%o su:
populations of Protamnio darwinii %ere aout to diver!e from each other0 over a very lon! time
and in a very i! %ay1 But they couldn(t diver!e if they kept on interreedin! %ith each other1 The
t%o !ene pools %ould continually flood each other %ith !enes1 So any tendency to diver!e %ould
e nipped in the ud efore it could !et !oin!0 s%amped y !ene flo% from the other population1
What actually happened at this epic partin! of the %ays0 noody kno%s1 #t happened a very
lon! time a!o0 and %e have no idea %here1 But modern evolutionary theory %ould confidently
reconstruct somethin! like the follo%in! history1 The t%o su:populations of Protamnio darwinii
someho% ecame separated from each other0 most likely y a !eo!raphical arrier such as a strip of
sea separatin! t%o islands0 or separatin! an island from a mainland1 #t could have een a mountain
ran!e that separated t%o valleys0 or a river separatin! t%o forestsA t%o (islands( in the !eneral sense #
defined1 &ll that matters is that the t%o populations %ere isolated from one another for lon! enou!h
so that0 %hen time and chance eventually reunited them0 they found they had diver!ed so much that
they couldn(t interreed any more1 )o% lon! is lon! enou!hB Well0 if they %ere suHect to stron!
and contrastin! selection pressures0 it could e as little as a fe% centuries0 or even less1 ,or
example0 an island mi!ht lack a voracious predator that pro%led the mainland1 Or the island
population mi!ht have shifted from an insectivorous to a ve!etarian diet0 like the &driatic liFards of
Chapter =1 Once a!ain0 %e can(t kno% the details of ho% Protamnio darwinii split0 and %e don(t
need to1 The evidence from modern animals !ives us every reason to think that somethin! like the
story # have Hust told is %hat happened in the past0 for every one of the diver!ences et%een the
ancestry of any animal and any other1
Even if conditions on either side of the arrier are identical0 t%o !eo!raphically separated
!ene pools of the same species %ill eventually drift apart from one another0 to the point %here they
can no lon!er interreed %hen the !eo!raphical isolation eventually comes to an end1 Random
chan!es in the t%o !ene pools %ill !radually uild up0 to the point %here0 if a male and a female
from the t%o sides meet0 their !enomes %ill e too different to comine to make a fertile offsprin!1
Whether y random drift alone0 or %ith the aid of differential natural selection0 once the t%o !ene
pools have reached the point %here they no lon!er need the !eo!raphical isolation to stay
!enetically separate0 %e call them t%o different species1 #n our hypothetical case0 perhaps the island
population chan!ed more than the mainland population0 ecause of the lack of predators and the
s%itch to a more ve!etarian diet1 So0 a Foolo!ist of the time mi!ht have reco!niFed that the island
population had ecome a ne% species and !iven it a ne% name0 say Protamnio sa"rops0 %hile the
old name0 Protamnio darwinii0 mi!ht have continued to serve for the mainland population1 #n our
hypothetical scenario0 perhaps it %as the island population that %as destined to !ive rise to the
sauropsid reptiles 7that(s everythin! %e call reptiles today plus irds80 %hile the mainland
population eventually !ave rise to the mammals1
Once a!ain0 # must stress0 the details of my little story are pure fiction1 #t could eEually %ell
have een the island population that !ave rise to the mammals1 The (island( could have een an oasis
surrounded y desert0 rather than land surrounded y %ater1 &nd of course %e haven(t the faintest
idea %hereaouts on the Earth(s surface this !reat divide took place : indeed0 the %orld map %ould
have looked so different that the Euestion scarcely means anythin!1 What is not fiction is the maHor
lessonA most0 if not all0 of the millions of evolutionary diver!ences that have populated the Earth
%ith such luxuriant diversity e!an %ith the chance separation of t%o su:populations of a species0
often0 thou!h not al%ays0 on either side of a !eo!raphical arrier such as a sea0 a river0 a mountain
ran!e or a desert valley1 Biolo!ists use the %ord (speciation( for the splittin! of a species into t%o
dau!hter species1 "ost iolo!ists %ill tell you that !eo!raphical isolation is the normal prelude to
speciation0 althou!h some0 especially entomolo!ists0 may chime in %ith the reservation that
(sympatric speciation( can also e important1 Sympatric speciation0 too0 reEuires some kind of
initial0 incidental separation to !et the all rollin!0 ut it is somethin! other than !eo!raphic
separation1 #t could e a local chan!e in microclimate1 # %on(t !o into the details0 ut %ill Hust say
that sympatric speciation seems to e especially important for insects1 *evertheless0 for simplicity(s
sake0 # shall in the rest of this chapter assume that the initial separation that precedes speciation is
normally !eo!raphical1 6ou may rememer that0 in Chapter 3(s treatment of domestic do! reeds0 #
likened the effect of the rules imposed y pedi!ree reeders to the creation of (virtual islands(1
(O*E "#G)T RE&--6 ,&*C6 1 1 1(
)o%0 then0 do t%o populations of a species find themselves on opposite sides of a
!eo!raphical arrierB Sometimes the arrier itself is the novelty1 &n earthEuake opens up an
impassale !or!e0 or chan!es the course of a river0 and a species that had een a sin!le reedin!
population finds itself severed in t%o1 "ore usually0 the arrier %as there all alon!0 and it is the
animals themselves that cross it0 in a rare freak event1 #t has to e rare0 other%ise it doesn(t deserve
to e called a arrier at all1 Before > Octoer 299= there %ere no memers of the species &$"ana
i$"ana on the Cariean island of &n!uilla1 On that date0 a population of these lar!e liFards
suddenly appeared on the eastern side of the island1 ,ortuitously0 they %ere actually seen arrivin!1
They %ere clin!in! to a mat of drift%ood and uprooted trees0 some more than 45 feet lon!0 that had
drifted from a nei!hourin! island0 proaly Guadeloupe 2<5 miles a%ay1 The previous month t%o
hurricanes0 -uis on >:= Septemer0 and "arilyn t%o %eeks later0 had ripped throu!h the area and
could easily have uprooted the trees0 complete %ith i!uanas0 %hich haitually spend time up trees1
The ne% population on &n!uilla %as still !oin! stron! in 299;0 and 'r Ellen Censky0 %ho led the
ori!inal study0 informs me that they are flourishin! to this day0 seemin!ly even more so than the
other species of i!uana that lived on &n!uilla efore the ne% invaders arrived1
The point aout such freak dispersal events is that they must e common enou!h to account
for speciation0 ut not too common1 #f they %ere too common : if0 say0 i!uanas drifted from
Guadeloupe to &n!uilla every year : the incipiently speciatin! population on &n!uilla %ould e
continually s%amped y incomin! !ene flo% and therefore could not diver!e from the Guadeloupe
population1 By the %ay0 please don(t e misled y my use of a phrase like (must e common
enou!h(1 #t could e misunderstood to mean that steps of some kind %ere taken to ensure that the
islands %ere Hust the ri!ht distance apart to facilitate speciationK Of course that puts the cart efore
the horse1 #t is rather that0 %herever there happen to e islands 7islands in the road sense0 as
al%ays8 spaced out at an appropriate distance to facilitate speciation0 there speciation %ill occur1
&nd the appropriate distance %ill depend on ho% easy it is for the animals concerned to travel1 The
2<5 miles that separate Guadeloupe from &n!uilla %ould e child(s play to any stron! flyin! ird
such as a petrel1 But even a sea crossin! of a fe% hundred yards mi!ht e difficult enou!h to
mid%ife a ne% species of0 say0 fro!s or %in!less insects1
The Galapa!os archipela!o is separated from the mainland of South &merica y aout <55
miles of open %ater0 nearly four times as far as those i!uanas sailed on their uprooted raft to
&n!uilla1 The islands are all volcanic0 and youn! y !eolo!ical standards1 *one of them has ever
een connected to any mainland1 The entire fauna and flora of the islands must have travelled there0
presumaly from mainland South &merica1 Even thou!h small irds can fly0 <55 miles is enou!h to
make a crossin! y finches a very rare event1 *ot so rare that it couldn(t happen0 ho%ever0 and there
are finches on Galapa!os0 %hose ancestors0 at some point in history0 %ere presumaly lo%n across0
perhaps y a freak storm1 These finches are all of a reco!niFaly South &merican type0 althou!h the
species themselves are uniEue to the Galapa!os islands1 -ook at 'ar%in(s map %hich # have
adopted for sentimental reasons and ecause he used the ma!nificently naval:soundin! En!lish
names for the islands0 rather than the modern Spanish names1 *otice that the <5:mile scale is aout
a tenth of the distance an animal %ould have had to travel to arrive on the archipela!o from the
mainland in the first place1 The islands themselves are only tens of miles from each other0 ut
hundreds of miles from the mainland1 What a %onderful recipe for speciation1 #t %ould e too
simple to say that the chance of ein! accidentally lo%n or rafted across a sea arrier to an island
is inversely proportional to the %idth of the arrier1 *evertheless0 there %ill clearly e some sort of
inverse correlation et%een distance and proaility of crossin!1 The difference et%een the
avera!e inter:island distance of a fe% tens of miles0 and the <55:mile distance to the mainland0 is so
lar!e that you %ould expect the archipela!o to e a po%erhouse of speciation1 &nd so it is0 as
'ar%in eventually realiFed0 althou!h not until after he had left the islands0 never to return1
Dar,in's "a# o$ the >ala#a'os islan%s ,ith 5n'lish na"es, no, sel%o" (se%
This disparity0 et%een tens of miles as the distance et%een islands %ithin the archipela!o0
and hundreds of miles as the distance of the %hole archipela!o from the mainland0 leads the
evolutionist to expect that the different islands mi!ht house species that are pretty similar to each
other ut more different from their counterparts on the mainland1 &nd that is exactly %hat %e do
find1 'ar%in himself put it %ell0 comin! tantaliFin!ly close to evolutionary lan!ua!e0 even efore
he had properly formulated his ideas1 # have placed the key clause in italics0 and shall repeat it
throu!hout this chapter in different contexts1
Seein! this !radation and diversity of structure in one small0 intimately related !roup of
irds0 one mi$ht really fancy that from an ori$inal pa"city of birds in this archipela$o* one species
had been taken and modified for different ends4 #n a like manner it mi!ht e fancied that a ird0
ori!inally a uFFard0 had een induced here to undertake the office of the carrion:feedin! Polyori
of the &merican continent1
The last sentence is a reference to the Galapa!os ha%k0 B"teo $alapa$oensis0 another
species that is found only on Galapa!os0 ut %hich some%hat resemles species on the mainland0
especially B"teo swainsoni0 %hich annually mi!rates et%een *orth and South &merica and could
%ell have een lo%n off course on one or t%o freak occasions1 *o%adays0 %e should refer to the
Galapa!os ha%k and the fli!htless cormorant as (endemic( to the islands0 meanin! that this is the
only place %here they are found1 'ar%in himself0 %ho had not yet fully emraced evolution0 used
the then current phrase (aori!inal creations(0 %hich meant that God had created them here and
no%here else1 )e used the same phrase of the !iant tortoises0 %hich then aounded on all the
islands0 and also of the t%o species of i!uana0 the Galapa!os land i!uana and the Galapa!os marine
i!uana1 The marine i!uanas are truly remarkale creatures0 Euite different from anythin! seen
any%here else in the %orld1 They dive to the sea ottom and !raFe sea%eed0 %hich seems to e their
only food1 They are !raceful s%immers0 althou!h not0 in 'ar%in(s outspoken vie%0 eautiful to look
#t is a hideous lookin! creature0 of a dirty lack colour0 stupid0L and slu!!ish in its
movements1 The usual len!th of a full:!ro%n one is aout a yard0 ut there are some even four feet
lon! 1 1 1 their tails are flattened side%ays0 and all four feet partially %eed 1 1 1 When in the %ater
this liFard s%ims %ith perfect ease and Euickness0 y a serpentine movement of its ody and
flattened tail : the le!s ein! motionless and closely collapsed on its sides1
Since marine i!uanas are so !ood at s%immin!0 it mi!ht e supposed that they0 rather than
the land i!uanas0 made the lon! crossin! from the mainland and suseEuently speciated0 in the
archipela!o0 to !ive rise to the land i!uana1 This is almost certainly not the case0 ho%ever1 The
Galapa!os land i!uana is not !reatly different from i!uanas still livin! on the mainland0 %hereas the
marine i!uanas are uniEue to the Galapa!os archipela!o1 *o liFard %ith the same marine haits has
ever een found else%here in the %orld1 We are no%adays confident that it %as the land i!uana that
ori!inally arrived from the South &merican mainland0 perhaps carted on drift%ood like the modern
ones from Guadeloupe that %ere lo%n to &n!uilla1 On Galapa!os0 they suseEuently speciated to
!ive rise to the marine i!uana1 &nd it %as almost certainly the !eo!raphical isolation permitted y
the spaced:out pattern of the islands that made possile the initial separation et%een the ancestral
land i!uanas and the ne%ly speciatin! marine i!uanas1 Presumaly some land i!uanas %ere
accidentally rafted across to a hitherto i!uana:free island0 and there adopted a marine hait0 free
from contamination y !enes flo%in! in from the land i!uanas on the ori!inal island1 "uch later0
they spread to other islands0 eventually returnin! to the island from %hich their land ancestors had
ori!inally hailed1 By no% they could no lon!er interreed %ith them0 and their !enetically inherited
marine haits %ere safe from contamination y land i!uana !enes1
#n example after example0 'ar%in noticed the same thin!1 The animals and plants of each
island of Galapa!os are lar!ely endemic to the archipela!o 7(aori!inal creations(80 ut they are also
for the most part uniEue0 in detail0 from island to island1 )e %as especially impressed %ith the
plants in this respectA
)ence %e have the truly %onderful fact0 that in .ames #sland MSantia!oN0 of the thirty:ei!ht
Galapa!eian plants0 or those found in no other part of the %orld0 thirty are exclusively confined to
this one islandJ and in &lemarle #sland M#saelaN0 of the t%enty:six aori!inal Galapa!eian plants0
t%enty:t%o are confined to this one island0 that is0 only four are at present kno%n to !ro% in the
other islands of the archipela!oJ and so on 1 1 1 %ith the plants from Chatham MSan CristoalN and
Charles M,loreanaN #slands1
)e noticed the same thin! %ith the distriution of mockin!irds over the islands1
"y attention %as first thorou!hly aroused0 y comparin! to!ether the numerous specimens0
shot y myself and several other parties on oard0 of the mockin!:thrushes0 %hen0 to my
astonishment0 # discovered that all those from Charles #sland elon!ed to one species 7%im"s
trifasciat"s8J all from &lemarle #sland to %4 par"l"sJ and all from .ames and Chatham #slands
7et%een %hich t%o other islands are situated0 as connectin! links8 elon!ed to %4 melanotis1
So it is0 all over the %orld1 The fauna and flora of a particular re!ion are Hust %hat %e should
expect if0 to Euote 'ar%in on the finches that no% ear his name0 (one species had een taken and
modified for different ends(1
The +ice:Governor of the Galapa!os #slands0 "r -a%son0 intri!ued 'ar%in y informin!
that the tortoises differed from the different islands0 and that he himself could %ith certainty
tell from %hich island any one %as rou!ht1 # did not for some time pay sufficient attention to this
statement0 and # had already partially min!led to!ether the collections from t%o of the islands1 #
never dreamed that islands0 aout fifty or sixty miles apart0 and most of them in si!ht of each other0
formed of precisely the same rocks0 placed under a Euite similar climate0 risin! to a nearly eEual
hei!ht0 %ould have een differently tenanted1
&ll the Galapa!os !iant tortoises are similar to a particular mainland species of land tortoise0
Geochelone chilensis0 %hich is smaller than any of them1 &t some point durin! the fe% million
years that the islands have existed0 one or a fe% of these mainland tortoises inadvertently fell in the
sea and floated across1 )o% could they have survived the lon! and doutless arduous crossin!B
Surely most of them didn(t1 But it %ould have only taken one female to do the trick1 &nd tortoises
are astonishin!ly %ell:eEuipped to survive the crossin!1
The early %halers took thousands of !iant tortoises from the Galapa!os islands a%ay in their
ships for food1 To keep the meat fresh0 the tortoises %ere not killed until needed0 ut they %ere not
fed or %atered %hile %aitin! to e utchered1 They %ere simply turned on their acks0 sometimes
stacked several deep0 so they couldn(t %alk a%ay1 # tell the story not in order to horrify 7althou!h #
have to say that such araric cruelty does horrify me80 ut to make a point1 Tortoises can survive
for %eeks %ithout food or fresh %ater0 easily lon! enou!h to float in the )umoldt Current from
South &merica to the Galapa!os archipela!o1 &nd tortoises do float1
)avin! reached and multiplied upon their first Galapa!os island0 the tortoises %ould %ith
comparative ease : a!ain accidentally : have island:hopped the much shorter distances to the rest of
the archipela!o y the same means1 &nd they did %hat many animals do %hen they arrive on an
islandA they evolved to ecome lar!er1 This is the lon!:noticed phenomenon of island !i!antism
7confusin!ly0 there is an eEually %ell:kno%n phenomenon of island d%arfism81L
#f the tortoises had follo%ed the pattern of 'ar%in(s famous finches0 they %ould have
evolved a different species on each of the islands1 Then0 after suseEuent accidental driftin!s from
island to island0 they %ould have een unale to interreed 7that(s the definition0 rememer0 of a
separate species8 and %ould have een free to evolve a different %ay of life uncontaminated y
!enetic s%ampin!1
6ou could say that the different species( incompatile matin! haits and preferences
constitute a kind of !enetic sustitute for the !eo!raphic isolation of separate islands1 Thou!h they
overlap !eo!raphically0 they are no% isolated on separate (islands( of matin! exclusivity1 So they
can diver!e yet further1 The -ar!e0 the "edium and the Small Ground ,inch ori!inally diver!ed on
different islandsJ the three species no% coexist on most of the Galapa!os islands0 never
interreedin! and each specialiFin! in a different kind of seed diet1
The tortoises did somethin! similar0 evolvin! distinctive shell shapes on the different
islands1 The species on the lar!er islands have hi!h domes1 Those on the smaller islands have
saddle:shaped shells %ith a hi!h:lipped %indo% at the front for the head1 The reason for this seems
to e that the lar!e islands are %et enou!h to !ro% !rass0 and the tortoises there are !raFers1 The
smaller islands are mostly too dry for !rass0 and the tortoises resort to ro%sin! on cactuses1 The
hi!h:lipped saddle shell allo%s the neck to reach up to the cactuses %hich0 for their part0 !ro%
hi!her in an evolutionary arms race a!ainst the ro%sin! tortoises1
The tortoise story adds to the finch model the further complication that0 for tortoises0
volcanoes are islands %ithin islands1 +olcanoes provide hi!h0 cool0 damp0 !reen oases0 surrounded
at lo% altitude y dry lava fields %hich0 for a !raFin! !iant tortoise0 constitute hostile deserts1 Each
of the smaller islands has a sin!le lar!e volcano and its o%n sin!le species 7or su:species8 of !iant
tortoise 7except those fe% islands that have none at all81 The i! island of #saela 7(&lemarle( to
'ar%in8 consists of a strin! of five maHor volcanoes0 and each volcano has its o%n species 7or su:
species8 of tortoise1 Truly0 #saela is an archipela!o %ithin an archipela!oA a system of islands
%ithin an island1 &nd the principle of islands in the literal !eo!raphical sense0 settin! the sta!e for
the evolution of islands in the metaphorical !enetic sense of species0 has never een more ele!antly
demonstrated than here in the archipela!o of 'ar%in(s lest youth1L
#slands don(t come much more isolated than St )elena0 a sin!le volcano in the South
&tlantic some 20355 miles from the coast of &frica1 #t has aout 255 endemic plants 7the youn!
'ar%in %ould have called them (aori!inal creations( and the older 'ar%in %ould have said they
evolved there81 &mon! these are 7or %ere0 for some of them are no% extinct8 forest trees elon!in!
to the daisy family1
These trees resemle in hait trees on the &frican mainland to %hich they are not closely
related1 The mainland plants to %hich they are related are hers or small shrus1 What must have
happened is that a fe% seeds of small hers or shrus chanced across the thousandmile !ap from
&frica0 settled on St )elena and0 ecause the niche of forest trees %as unfilled0 evolved lar!er and
more %oody trunks until they ecame proper trees1 Similar tree:like daisies have evolved
independently on the Galapa!os archipela!o1 #t is the same pattern on islands the %orld over1
Forest trees on St Helena
Each of the !reat &frican lakes has its o%n uniEue fish fauna0 dominated y the !roup called
cichlids1 The cichlid faunas of -ake +ictoria0 -ake Tan!anyika and -ake "ala%i0 each several
hundred species stron!0 are completely distinct from each other1 They have evidently evolved
separately in the three lakes0 %hich makes it all the more fascinatin! that they have conver!ed on
the same ran!e of (trades( in all three1 #n each lake0 it looks as thou!h one or t%o founder species
someho% made their %ay in0 perhaps from rivers0 in the first place1 &nd in each lake these founders
then speciated and speciated a!ain0 to populate the lake %ith the hundreds of species that %e see
today1 )o%0 %ithin the confines of a lake0 did the uddin! species achieve the initial !eo!raphical
isolation that enaled them to split apartB
When introducin! islands0 # explained that0 from a fish(s point of vie%0 a lake surrounded y
land is an island1 Sli!htly less oviously0 even an island in the conventional sense of land
surrounded y %ater can e an (island( for a fish0 especially a fish that lives only in shallo% %ater1 #n
the sea0 think of a coral:reef fish0 %hich never ventures into deep %ater1 ,rom its point of vie%0 the
shallo% frin!e of a coral island is an (island(0 and the Great Barrier Reef is an archipela!o1
Somethin! similar can happen even in a lake1 Within a lake0 especially a lar!e one0 a rocky outcrop
can e an (island( for a fish %hose haits confine it to shallo% %ater1 This is almost certainly ho% at
least some of the cichlids in the &frican !reat lakes achieved their initial isolation1 "ost individuals
%ere confined to shallo% %ater around islands0 or in ays and inlets1 This achieved partial isolation
from other such pockets of shallo% %ater0 linked y occasional traversin!s of the deeper %ater
et%een them to form the %atery eEuivalent of a Galapa!os:like (archipela!o(1
There(s !ood evidence 7for example from sediment core samples8 that the level of -ake
"ala%i 7it %as called -ake *yasa %hen # spent my first ucket:and:spade holidays on its sandy
eaches8 rises and falls dramatically over the centuries0 and reached a lo% point in the ei!hteenth
century0 more than 255 metres lo%er than the present level1 "any of its islands %ere not islands at
all durin! that time0 ut hills on the land around the then smaller lake1 When the lake level rose0 in
the nineteenth and t%entieth centuries0 the hills ecame islands0 ran!es of hills ecame
archipela!oes0 and the process of speciation took off amon! the cichlids that live in shallo% %ater0
kno%n locally as "una1 (&lmost every rocky outcrop and island has a uniEue "una fauna0 %ith
endless colour forms and species1 &s many of these islands and outcrops %ere dry land %ithin the
last 355:455 years0 the estalishment of the faunas has taken place %ithin that time1(
Such rapid speciation is somethin! the cichlid fishes are extremely !ood at1 -ake "ala%i
and -ake Tan!anyika are old0 ut -ake +ictoria is extremely youn!1 The lake asin %as formed
only aout >550555 years a!o0 and it has dried up several times since then0 most recently aout
2@0555 years a!o1 This seems to mean that its endemic fauna of >=5 or so species of cichlid fishes
have all evolved over a timescale of centuries0 not the millions of years that %e usually associate
%ith evolutionary diver!ence on this !rand scale1 The cichlids of &frica(s lakes impress us mi!htily
%ith %hat evolution can do in a short space of time1 They almost Eualified for inclusion in the
(efore our very eyes( chapter1
The %oods and forests of &ustralia are dominated y trees of a sin!le !enus0 E"calypt"s0
and there are more than @55 species of them0 fillin! a hu!e ran!e of niches1 Once a!ain0 'ar%in(s
dictum aout finches can e cooptedA one could almost ima!ine that one species of eucalypt had
een (taken and modified for different ends(1 &nd0 alon! parallel lines0 an even more famous
example is the &ustralian mammal fauna1 #n &ustralia there are0 or %ere until recent extinctions
possily caused y the arrival of aori!inal people0 the ecolo!ical eEuivalents of %olves0 cats0
raits0 moles0 shre%s0 lions0 flyin! sEuirrels and many others1 6et they are marsupials0 Euite
different from the %olves0 cats0 raits0 moles0 shre%s0 lions and flyin! sEuirrels %ith %hich %e are
familiar in the rest of the %orld0 the so:called placental mammals1 The &ustralian eEuivalents are all
descended from Hust a fe%0 or even one0 ancestral marsupial species0 (taken and modified for
different ends(1 This eautiful marsupial fauna has also produced creatures for %hich it is harder to
find a counterpart outside &ustralia1 The many species of kan!aroo mostly fill antelope:like niches
7or monkey or lemur:like niches in the case of the tree kan!aroos8 ut !et aout y hoppin! rather
than !allopin!1 They ran!e from the lar!e red kan!aroo 7and some even lar!er extinct ones0
includin! a fearsome0 oundin! carnivore8 to the small %allaies and tree kan!aroos1 There %ere
!iant0 rhinoceros:siFed marsupials0 'iprotodonts0 related to modern %omats ut 4 yards lon!0 <
feet tall at the shoulder0 and %ei!hin! 3 tons1 # shall return to the marsupials of &ustralia in the next
#t is almost too ridiculous to mention it0 ut #(m afraid # have to ecause of the more than >5
per cent of the &merican population %ho0 as # lamented in Chapter 20 accept the Bile literallyA
think %hat the !eo!raphical distriution of animals should look like if they(d all dispersed from
*oah(s &rk1 Shouldn(t there e some sort of la% of decreasin! species diversity as %e move a%ay
from an epicentre : perhaps "ount &raratB # don(t need to tell you that that is not %hat %e see1
Why %ould all those marsupials : ran!in! from tiny pouched mice throu!h koalas and
ilys to !iant kan!aroos and 'iprotodonts : %hy %ould all those marsupials0 ut no placentals at
all0 have mi!rated en masse from "ount &rarat to &ustraliaB Which route did they takeB &nd %hy
did not a sin!le memer of their stra!!lin! caravan pause on the %ay0 and settle : in #ndia0 perhaps0
or China0 or some haven alon! the Great Silk RoadB Why did the entire order Edentata 7all t%enty
species of armadillo0 includin! the extinct !iant armadillo0 all six species of sloth0 includin! extinct
!iant sloths0 and all four species of anteater8 troop off unerrin!ly for South &merica0 leavin! not a
rack ehind0 leavin! no hide nor hair nor armour plate of settlers some%here alon! the %ayB Why
%ere they Hoined y the entire infraorder of caviomorph rodents0 includin! !uinea pi!s0 a!outis0
pacas0 maras0 capyaras0 chinchillas and lots of others0 a lar!e !roup of characteristically South
&merican rodents0 found no%here elseB Why did an entire su:order of monkeys0 the platyrrhine
monkeys0 end up in South &merica and no%here elseB Shouldn(t at least a fe% of them have Hoined
the rest of the monkeys0 the catarrhines0 in &sia or &fricaB &nd shouldn(t at least one species of
catarrhine have found itself in the *e% World0 alon! %ith the platyrrhinesB Why did all the
pen!uins undertake the lon! %addle south to the &ntarctic0 not a sin!le one to the eEually
hospitale &rcticB
&n ancestral lemur0 a!ain very possily Hust a sin!le species0 found itself in "ada!ascar1
*o% there are thirty:seven species of lemur 7plus some extinct ones81 They ran!e in siFe from the
py!my mouse lemur0 smaller than a hamster0 to a !iant lemur0 lar!er than a !orilla and resemlin! a
ear0 %hich %ent extinct Euite recently1 &nd they are all0 every last one of them0 in "ada!ascar1
There are no lemurs any%here else in the %orld0 and there are no monkeys in "ada!ascar1 )o% on
Earth do the >5 per cent history:deniers think this state of affairs came aoutB 'id all thirty:seven
and more species of lemur troop in a ody do%n *oah(s !an!plank and hi!htail it 7literally in the
case of the rin!tail8 for "ada!ascar0 leavin! not a sin!le stra!!ler y the %ayside0 any%here
throu!hout the len!th and readth of &fricaB
Once a!ain0 # am sorry to take a sled!ehammer to so small and fra!ile a nut0 ut # have to do
so ecause more than >5 per cent of the &merican people elieve literally in the story of *oah(s
&rk1 We should e ale to i!nore them and !et on %ith our science0 ut %e can(t afford to ecause
they control school oards0 they home:school their children to deprive them of access to proper
science teachers0 and they include many memers of the $nited States Con!ress0 some state
!overnors and even presidential and vice:presidential candidates1 They have the money and the
po%er to uild institutions0 universities0 even a museum %here children ride life:siFe mechanical
models of dinosaurs0 %hich0 they are solemnly told0 coexisted %ith humans1 &nd0 as recent polls
have sho%n0 Britain is not far ehind 7or should that read (ahead(B80 alon! %ith parts of Europe and
most of the #slamic %orld1
Even if %e leave "ount &rarat to one sideJ even if %e refrain from lampoonin! those %ho
take the *oah(s &rk myth literally0 similar prolems apply to any theory of the separate creation of
species1 Why %ould an all:po%erful creator decide to plant his carefully crafted species on islands
and continents in exactly the appropriate pattern to su!!est0 irresistily0 that they had evolved and
dispersed from the site of their evolutionB Why %ould he put lemurs in "ada!ascar and no%here
elseB Why put platyrrhine monkeys in South &merica only0 and catarrhine monkeys in &frica and
&sia onlyB Why no mammals in *e% Iealand0 except ats %ho could fly thereB Why do the
animals in island chains most closely resemle those on nei!hourin! islands0 and %hy do they
nearly al%ays resemle : less stron!ly ut still unmistakaly : those on the nearest continent or
lar!e islandB Why %ould the creator put only marsupial mammals in &ustralia0 a!ain except ats
%ho could fly there0 and those %ho could arrive in man:made canoesB The fact is that0 if %e survey
every continent and every island0 every lake and every river0 every mountaintop and every &lpine
valley0 every forest and every desert0 the only %ay to make sense of the distriution of animals and
plants is0 yet a!ain0 to follo% 'ar%in(s insi!ht aout the Galapa!os finchesA (One mi!ht really fancy
that from an ori!inal paucity 1 1 1 one species had een taken and modified for different ends1(
'ar%in %as fascinated y islands0 and he tramped the len!th and readth of a !ood fe%
durin! the voya!e of the Bea$le1 )e even %orked out the surprisin! truth aout ho% islands of one
maHor class0 those uilt y the animals called corals0 are formed1 'ar%in later came to reco!niFe the
crucial importance of islands and archipela!oes for his theory0 and he did several experiments to
settle Euestions aout the theory of !eo!raphical isolation as a prelude to speciation 7he didn(t use
the %ord81 ,or example0 in a numer of experiments he kept seeds in sea %ater for lon! periods0 and
demonstrated that some retained the po%er to !erminate even after immersion for lon! enou!h to
have drifted from continents to nei!hourin! islands1 ,ro!spa%n0 on the other hand0 he found to e
immediately killed y sea %ater0 and he made !ood use of this to explain a si!nal fact aout the
!eo!raphical distriution of fro!sA
With respect to the asence of %hole orders on oceanic islands0 Bory St1 +incent lon! a!o
remarked that Batrachians 7fro!s0 toads0 ne%ts8 have never een found on any of the many islands
%ith %hich the !reat oceans are studded1 # have taken pains to verify this assertion0 and # have found
it strictly true1 # have0 ho%ever0 een assured that a fro! exists on the mountains of the !reat island
of *e% IealandJ ut # suspect that this exception 7if the information e correct8 may e explained
throu!h !lacial a!ency1 This !eneral asence of fro!s0 toads0 and ne%ts on so many oceanic islands
cannot e accounted for y their physical conditionsJ indeed it seems that islands are peculiarly %ell
fitted for these animalsJ for fro!s have een introduced into "adeira0 the &Fores0 and "auritius0
and have multiplied so as to ecome a nuisance1 But as these animals and their spa%n are kno%n to
e immediately killed y sea:%ater0 on my vie% %e can see that there %ould e !reat difficulty in
their transportal across the sea0 and therefore %hy they do not exist on any oceanic island1 But %hy0
on the theory of creation0 they should not have een created there0 it %ould e very difficult to
'ar%in %as %ell a%are of the si!nificance of the !eo!raphical distriution of species for his
theory of evolution1 )e noted that most of the facts could e accounted for if %e assume that
animals and plants have evolved1 ,rom this0 %e should expect : and %e find : that modern animals
tend to live on the same continent as fossils that could plausily e their ancestors0 or close to their
ancestors1 We should expect0 and %e find0 that animals share the same continent %ith species that
resemle them1 )ere is 'ar%in on the suHect0 payin! special attention to the animals of South
&merica that he kne% so %ellA
the naturalist in travellin!0 for instance0 from north to south never fails to e struck y the
manner in %hich successive !roups of ein!s0 specifically distinct0 yet clearly related0 replace each
other1 )e hears from closely allied0 yet distinct kinds of irds0 notes nearly similar0 and sees their
nests similarly constructed0 ut not Euite alike0 %ith e!!s coloured in nearly the same manner1 The
plains near the Straits of "a!ellan are inhaited y one species of Rhea 7&merican ostrich80 and
north%ard the plains of -a Plata y another species of the same !enusJ and not y a true ostrich or
emeu0 like those found in &frica and &ustralia under the same latitude1 On these same plains of -a
Plata0 %e see the a!outi and iFcacha0 animals havin! nearly the same haits as our hares and
raits0 1 1 1 ut they plainly display an &merican type of structure1 We ascend the lofty peaks of the
Cordillera and %e find an alpine species of iFcachaJ %e look to the %aters0 and %e do not find the
eaver or musk:rat0 ut the coypu and capyara0 rodents of the &merican type1
This is mostly common sense0 and 'ar%in %as ale to account for an enormous ran!e of
oservations y means of it1 But there are certain facts aout the !eo!raphical distriution of
animals and plants0 and the distriution of rocks0 that need a different kind of explanationA one that
is anythin! ut common sense0 and %hich %ould have sta!!ered and enthralled 'ar%in0 if only he
had kno%n aout it1
'#' T)E E&RT) "O+EB
Everyody in 'ar%in(s time thou!ht that the map of the %orld %as pretty much a constant1
Some of 'ar%in(s contemporaries did countenance the possiility of lar!e land rid!es0 no%
sumer!ed0 to explain0 for example0 the similarities et%een the floras of South &merica and
&frica1 'ar%in himself %as not !reatly enamoured of the land rid!e idea0 ut he surely %ould have
exulted in the modern evidence that entire continents move over the face of the Earth1 This provides
y far the est explanation of certain maHor facts of animal and plant dispersion0 especially of
fossils1 ,or example0 there are similarities et%een the fossils of South &merica0 &frica0 &ntarctica0
"ada!ascar0 #ndia and &ustralia0 %hich no%adays %e explain y invokin! the once !reat southern
continent of Gond%ana0 unitin! all those modern lands1 Once a!ain0 our late:comin! detective is
forced to the conclusion that evolution is a fact1
The theory of (continental drift(0 as it used to e called0 %as first championed y the German
climatolo!ist &lfred We!ener 72;;5:294581 We!ener %as not the first to look at a map of the %orld
and notice that the shape of a continent or island often matches the coastline opposite as if the t%o
land masses %ere pieces of a Hi!sa% puFFle0 even %hen the opposite coastline is far a%ay1 #(m not
talkin! aout little local examples0 such as the #sle of Wi!ht(s neat dovetailin! into the )ampshire
coast0 almost as thou!h the Solent %asn(t there1 What We!ener and his predecessors noticed %as
that somethin! alon! the same lines seemed to e true of the %hole facin! sides of the !iant
continents of &frica and &merica1 The BraFilian coast looks tailor:cut to fit under the ul!e of West
&frica0 %hile the northern part of &frica(s ul!e is a nice fit to the *orth &merican coast from
,lorida to Canada1 *ot only do the shapes rou!hly matchA We!ener also pointed to matchin!
!eolo!ical formations up and do%n the east side of South &merica and correspondin! parts of the
%est side of &frica1 Sli!htly less clearly0 the %est coast of "ada!ascar forms a pretty !ood fit to the
east coast of &frica 7not the section of South &frican coast that is opposite it today ut the coast of
TanFania and Denya further north80 %hile the lon!0 strai!ht line of "ada!ascar(s east side is
comparale to the strai!ht ed!e of %estern #ndia1 We!ener also pointed out that the ancient fossils
to e found in &frica and South &merica %ere more similar than you %ould expect if the map of the
%orld had al%ays een the %ay it is today1 )o% could this e0 !iven the %idth of the South &tlantic
oceanB Were the t%o continents once much closer0 or even HoinedB The idea %as tantaliFin!0 ut
ahead of its time1 We!ener also noticed matchin!s et%een the fossils of "ada!ascar and #ndia1
&nd there are similarly tellin! affinities et%een the fossils of northern *orth &merica and of
Such oservations led We!ener to propose his darin!ly heretical hypothesis of continental
drift1 &ll the !reat continents of the %orld0 he su!!ested0 had once een Hoined up in a !i!antic
super:continent0 %hich he called Pan!aea1 Over an immense span of !eolo!ical time0 he proposed0
Pan!aea !radually dismemered itself to form the continents %e kno% today0 %hich then slo%ly
drifted to their present positions and have not finished driftin! yet1
One can almost hear We!ener(s sceptical contemporaries %onderin!0 to use the street:talk of
today0 %hat he had een smokin!1 6et %e no% kno% that he %as ri!ht1 Or almost ri!ht1 ,ar:si!hted
and ima!inative as We!ener %as0 # must make it clear that his hypothesis of continental drift %as
si!nificantly different from our modern theory of plate tectonics1 We!ener thou!ht that the
continents plou!hed throu!h the oceans like !i!antic ships0 not Euite floatin! in %ater like 'r
'olittle(s hollo% island of Popsipetl0 ut floatin! atop the semi:liEuid mantle of the planet1
Reasonaly enou!h0 other scientists erected fortresses of scepticism1 What titanic forces could
propel an oHect the siFe of South &merica or &frica for thousands of milesB # shall explain ho% the
modern theory of plate tectonics differs from We!ener(s theory efore comin! to its supportin!
Cartoon ins#ire% *y We'ener's 'ontinental %ri$t' theory
#n the theory of plate tectonics the %hole of the Earth(s surface0 includin! the ottoms of the
various oceans0 consists of a series of overlappin! rocky plates like a suit of armour1 The continents
that %e see are thickenin!s of the plates that rise aove sea level1 The !reater part of the area of
each plate lies under the sea1 $nlike We!ener(s continents0 the plates do not sail throu!h the sea0 or
plou!h throu!h the surface of the Earth0 they are the surface of the Earth1 'on(t think0 like
We!ener0 of the continents themselves as ein! Hi!sa%ed to!ether or ein! pulled apart from each
otherJ it isn(t like that1 Think of a plate0 instead0 as ein! continuously manufactured at a !ro%in!
ed!e0 in a remarkale process called sea:floor spreadin!0 %hich # shall explain in a moment1 &t
other ed!es0 a plate may e (suducted( under a nei!hourin! plate1 Or nei!hourin! plates may
slide alon!side one another1 The picture on colour pa!e 2@ sho%s a portion of the San &ndreas
,ault in California0 %hich is %here the ed!es of the Pacific plate and the *orth &merican plate
shear past each other1 The comination of sea:floor spreadin! and suduction means that there are
no !aps et%een plates1 The entire surface of the planet is covered in plates0 each one typically
disappearin! y suduction underneath a nei!hourin! plate on one side0 or slidin! past another
plate0 %hile it !ro%s out from a sea:floor spreadin! Fone else%here1
#t is inspirin! to think of the hu!e rift valley that must once have snaked its %ay do%n the
continent of Gond%ana et%een the future &frica and future South &merica1 *o dout it %as at
first dotted %ith lakes like the present rift valley of East &frica1 -ater0 it filled %ith sea %ater as
South &merica sheared a%ay %ith rendin! tectonic a!ony1 #ma!ine the vie% that !reeted some stout
dinosaurian CorteF as he !aFed across the lon!0 narro% straits at the slo%ly departin! (West
Gond%ana(1 We!ener %as ri!ht that the Hi!sa% complementarity of their shapes is no accident1 But
he %as %ron! to think of the continents as !i!antic rafts0 plou!hin! their %ay throu!h the seafilled
!aps et%een them1 South &merica and &frica0 and their continental shelves0 are ut the thickened
re!ions of t%o plates0 much of %hose rocky surfaces lies under the sea1 The plates constitute the
hard lithosphere : literally0 (sphere of rock( : %hich floats atop the hot0 semi:molten asthenosphere :
(sphere of %eakness(1 The asthenosphere is %eak in the sense that it is not ri!id and rittle like the
rocky plates of the lithosphere ut ehaves some%hat like a liEuidA yieldin!0 like putty or toffee0 if
not necessarily molten1 & little confusin!ly0 this distinction et%een t%o concentric spheres doesn(t
%holly correspond to the more familiar distinction 7ased on chemical composition rather than
physical stren!th8 et%een the (crust( and the (mantle(1
"ost plates consist of t%o distinct kinds of lithospheric rock1 The deep ocean ottoms are
covered in a rather uniform layer of very dense i!neous rock0 aout 25 kilometres thick1 This
i!neous layer is overlain y a superficial layer of sedimentary rock and mud1 & continent0 to repeat0
is the area of a plate visile aove sea level0 risin! to this hei!ht %here the plate is thickened y
additional layers of less dense rock1 The undersea parts of the plates are continuously ein! created
at their mar!ins : the eastern mar!in in the case of the South &merican plate0 the %estern mar!in in
the case of the &frican plate1 These t%o mar!ins comprise the mid:&tlantic rid!e0 %hich snakes its
%ay do%n the middle of the &tlantic from #celand 7%hich is0 indeed0 the only sustantial part of the
rid!e that reaches the surface8 to the far south1
Similar undersea rid!es are rollin! out other plates in other parts of the %orld 7see colour
pa!es 2;:2981 These undersea rid!es %ork like elon!ated fountains 7on the slo% timescale of
!eolo!y80 %ellin! up molten rock in the process # have already mentioned called sea:floor
spreadin!1 The spreadin! sea:floor rid!e in mid:&tlantic seems to push the &frican plate east%ards
and the South &merican plate %est%ards1 The ima!e of a pair of rolltop desks spreadin! in
diver!ent directions has een su!!ested0 and it conveys the idea0 provided %e rememer that it is all
happenin! on a timescale too slo% for humans to see1 #ndeed0 the speed at %hich South &merica
and &frica pull apart has een memoraly likened : so memoraly that it has ecome almost a
cliche : to the speed at %hich fin!ernails !ro%1 The fact that they are no% thousands of miles apart
is further testimony to the vast and unilical a!e of the Earth0 comparale to the evidence from
radioactivity %hich %e met in Chapter >1
# used the phrase (seems to push( Hust no%0 and # must hasten to acktrack1 #t is temptin! to
think of those up%ellin! (rolltop desks( as pushin! their respective continental plates alon! from
ehind1 This is unrealisticJ the scale is all %ron!1 Tectonic plates are much too massive to e pushed
from ehind y up%ellin! volcanic forces alon! a mid:ocean rid!e1 & s%immin! tadpole mi!ht as
%ell try to push a supertanker1 But no%0 here(s the key point1 The asthenosphere0 in its capacity as a
Euasi:liEuid0 has convection currents that extend throu!hout its %hole surface0 under the entire area
of the plates1 #n any one re!ion0 the asthenosphere is slo%ly movin! in a consistent direction0 and
then circlin! ack in the opposite direction do%n in its deeper layers1 The upper layer of
asthenosphere under the South &merican plate0 for example0 is movin! inexoraly %est%ard1 &nd0
%hile it is inconceivale that an up%ellin! (rolltop desk( could have the stren!th to push the %hole
South &merican plate efore it0 it is not at all inconceivale that a convection current inchin! its
%ay steadily in a consistent direction "nder the entire lower s"rface of a plate could carry its
(floatin!( continental urden alon! %ith it1 We aren(t talkin! tadpoles no%1 & supertanker in the
)umoldt Current0 %ith its en!ines s%itched off0 %ill indeed !o %ith the flo%1
That is0 in summary0 the modern theory of plate tectonics1 # must no% turn to the evidence
that it is true1 &ctually0 as is normal %ith estalished scientific facts0L there are lots of different
kinds of evidence0 ut # am only !oin! to talk aout the most strikin!ly ele!ant kind1 This is the
evidence from the a!es of the rocks0 and especially from the ma!netic stripes in them1 #t(s almost
too !ood to e true0 a perfect illustration of my (detective comin! late to the scene of the crime( and
ein! driven inexoraly to only one conclusion1 We even have somethin! that looks very like
fin!erprintsA !iant ma!netic fin!erprints in the rocks1
We shall accompany our metaphorical detective on a voya!e across the South &tlantic0 in a
custom:uilt sumarine capale of %ithstandin! the dauntin! pressures of the deep sea1 The
sumarine is eEuipped to drill do%n for rock samples0 throu!h the superficial sediments of the sea
ottom do%n to the volcanic rocks of the lithosphere itself0 and it also has an onoard laoratory for
datin! rock samples radiometrically 7see Chapter >81 The detective sets a course due east from the
BraFilian port of "aceio0 25 de!rees of latitude south of the eEuator1 )avin! travelled =5 kilometres
or so throu!h the shallo% %aters of the continental shelf 7%hich for present purposes counts as part
of South &merica8 %e atten do%n the hi!h:pressure hatches and dive 7%hat understatementK80 dive
do%n into the depths %here the only li!ht normally seen is the occasional spark of !reenish
luminescence from the !rotesEues that inhait this alien %orld1
When %e hit ottom at nearly 350555 feet 7full fathom 4055580 %e drill do%n to the volcanic
lithosphere and take a core sample of the rock1 The onoard radioactive datin! la !oes to %ork0
and reports a -o%er Cretaceous a!e0 aout 2>5 million years1 The sumarine !rinds on east%ards
alon! the tenth parallel0 takin! rock samples at freEuent intervals1 The a!e of each sample is
carefully measured and the detective pores over the datin!s0 lookin! for a pattern1 )e doesn(t have
to look far1 Even 'r Watson couldn(t miss it1 &s %e travel east alon! the !reat plains of the sea
ottom0 the rocks are plainly !ettin! youn!er and youn!er0 steadily youn!er1 &out @45 kilometres
into our Hourney0 the rock samples are of late Cretaceous a!e0 aout <= million years old0 %hich
happens to e %hen the last of the dinosaurs %ent extinct1 The trend to%ards youn!er and youn!er
rocks continues as %e approach the middle of the &tlantic and the sumarine(s searchli!hts start to
pick out the foothills of a !i!antic under%ater mountain ran!e1 This is the mid:&tlantic rid!e 7see
colour pa!es 2;:2980 %hich our sumarine must no% start to clim1 $p and up %e cra%l0 still takin!
rock samples0 and still noticin! that the rocks are !ettin! youn!er and youn!er1 By the time %e
reach the peaks of the rid!e0 the rocks are so youn! they mi!ht as %ell have only Hust %elled up as
fresh lava from volcanoes1 #ndeed0 that is pretty much %hat has happened1 &scension #sland is a
part of the mid:&tlantic rid!e %hich protruded aove sea level as a result of a recent series of
eruptions : %ell0 recentA maye < million years a!oJ that(s recent y the standards of the rocks %e
have een samplin! alon! our sumarine %ay1
We no% push on to%ards &frica0 over the other side of the rid!e0 do%n to the deep plains at
the ottom of the eastern &tlantic1 We continue to take rock samples and : you(ve !uessed it : the
rocks no% ecome steadily older as %e move to%ards &frica1 #t is the mirror ima!e of the pattern
%e noticed efore %e reached the mid:&tlantic rid!e1 The detective is in no dout of the
explanation1 The t%o plates are movin! apart as the sea floor spreads a%ay from the rid!e1 &ll the
ne% rock that is ein! added to the t%o diver!in! plates comes from the volcanic activity of the
rid!e itself0 and it is then carried a%ay0 in opposite directions0 on one or other of the !i!antic rolltop
desks that %e call the &frican plate and the South &merican plate1 The false colours in the pictures
on colour pa!es 2;:29 illustratin! this process denote the a!e of the rocks0 red ein! the youn!est1
6ou can see ho% eautifully the a!e profiles on the t%o sides of the mid:&tlantic rid!e mirror each
What an ele!ant storyK But it !ets etter1 The detective notices a more sutle pattern in the
rock samples as they are processed in the onoard laoratory1 The rock cores pulled up from the
deep lithosphere are sli!htly ma!netic0 like compass needles1 The phenomenon is %ell understood1
When molten rock solidifies0 the Earth(s ma!netic field ecomes imprinted into it0 in the form of a
polariFation of the fine crystals of %hich i!neous rock is made1 The crystals ehave like tiny froFen
compass needles0 locked into the direction they %ere pointin! in at the moment %hen the molten
lava solidified1 *o%0 it has lon! een kno%n that Earth(s ma!netic pole is not fixed ut %anders0
proaly ecause of slo%ly ooFin! currents in the mixture of molten iron and nickel in the planet(s
core1 &t present0 ma!netic north lies near Ellesmere #sland in northern Canada0 ut it %on(t stay
there1 To determine true north usin! a ma!netic compass0 sailors need to look up a correction factor0
and the correction factor chan!es from year to year as the planet(s ma!netic field fluctuates1
So lon! as our detective meticulously records the exact an!le at %hich his rocky cores sat
%hen he drilled them out0 the froFen ma!netic field in each core tells him the position of the Earth(s
ma!netic field on the day that the rock solidified out of lava1 &nd no% for the clincher1 #t happens
that0 at irre!ular intervals of tens or hundreds of thousands of years0 the Earth(s ma!netic field
completely reverses0 presumaly ecause of maHor shifts in the molten nickelG iron core1 What %as
ma!netic north flips over to a position near the true South Pole0 and %hat %as ma!netic south flips
to the north1 &nd of course the rocks pick up the current position of ma!netic north on the day that
they solidify from lava %ellin! up from the deep sea ottom1 Because the polariFation reverses
every fe% tens of thousands of years0 a ma!netometer can detect stripes runnin! alon! the edrockA
stripes in %hich the rock samples( ma!netic fields all point in one direction0 alternatin! %ith stripes
in %hich the ma!netic fields all point in the opposite direction1 Our detective colours them lack
and %hite on the map1 &nd %hen he looks at the stripes on the map like fin!erprints0 he notices an
unmistakale pattern1 &s %ith the false colour stripes denotin! the asolute a!e of the rocks0 the
ma!netic fin!erprint stripes on the %est side of the mid:&tlantic rid!e are an ele!ant mirror ima!e
of the stripes on the east side1 Exactly %hat you(d expect if the ma!netic polarity of the rock %as
laid do%n %hen the lava first solidified in the rid!e and then slo%ly moved out%ards from the
rid!e0 in opposite directions0 at a fixed and very slo% rate1 Elementary0 my dear Watson1 L
To revert to the terminolo!y of Chapter 20 the morphin! of We!ener(s hypothesis of
continental drift into the modern theory of plate tectonics is a textook example of the solidification
of a tentative hypothesis into a universally accepted theorum or fact1 Plate tectonic movements are
important in this chapter0 ecause %ithout them %e cannot fully understand the distriution of
animals and plants over the continents and islands of the %orld1 When # spoke of the initial
!eo!raphical arrier that separated t%o incipient species0 # proposed an earthEuake divertin! the
course of a river1 # could also have mentioned plate tectonic forces0 splittin! a continent in t%o and
ferryin! the t%o !i!antic fra!ments in opposite directions0 complete %ith animal and plant
passen!ers : the ark of the continents1
"ada!ascar and &frica %ere once part of the !reat southern continent of Gond%ana0
to!ether %ith South &merica0 &ntarctica0 #ndia and &ustralia1 Gond%ana e!an to reak up :
painfully slo%ly y the standards of our perception : aout 2<= million years a!o1 &t this point
"ada!ascar0 %hile still Hoined to #ndia0 &ustralia and &ntarctica as East Gond%ana0 pulled a%ay
from the eastern side of &frica1 &t aout the same time0 South &merica pulled a%ay from West
&frica in the other direction1 East Gond%ana itself roke up rather later0 and "ada!ascar finally
ecame separated from #ndia aout 95 million years a!o1 Each of the fra!ments of old Gond%ana
carried %ith it its car!o of animals and plants1 "ada!ascar %as a real (ark(0 and #ndia %as another1 #t
is proale0 for example0 that the ancestors of ostriches and elephant irds ori!inated in
"ada!ascarG#ndia %hen they %ere still united1 -ater they split1 Those that remained on the !iant raft
called "ada!ascar evolved to ecome the elephant irds0 %hile the ancestors of ostriches sailed off
on the !ood ship #ndia and suseEuently : %hen #ndia collided %ith &sia and raised the )imalayas :
%ere lierated on to the mainland of &sia0 %hence they eventually found their %ay to &frica0 their
main stampin! !round today 7yes0 the males really do stamp their feet0 to impress females81
Elephant irds0 alas0 %e no lon!er see 7or hear0 more(s the pity0 for if they stamped the very !round
must have shaken81 ,ar more massive than the lar!est ostriches0 these "ada!ascar !iants are the
proale ori!in of the le!endary (roc(0 %hich features in the Second +oya!e of Sinad the Sailor1
&lthou!h lar!e enou!h for a man to have ridden0 they had no %in!s0 so could never have carried
Sinad aloft as advertised1 L
*ot only does the no% solidly estalished theory of plate tectonics account for numerous
facts aout the distriution of fossils and livin! creatures0 it also provides yet more evidence of the
extreme antiEuity of the Earth1 #t ou!ht0 therefore0 to e a maHor thorn in the side of creationists0 at
least creationists of the (youn! Earth( persuasion1 )o% do they cope %ith itB +ery %eirdly indeed1
They don(t deny the shiftin! of the continents0 ut they think it all happened at hi!h speed very
recently0 at the time of *oah(s flood1 L 6ou(d think that0 since they are conspicuously happy to
discount evidence that doesn(t suit them in the case of the massive Euantity and ran!e of evidence
for the fact of evolution0 they(d pull the same trick %ith the evidence for plate tectonics too1 But noA
oddly0 they accept as a fact that South &merica once fitted snu!ly into &frica1 They seem to re!ard
the evidence for this as conclusive0 even thou!h the evidence for the fact of evolution is0 if
anythin!0 even stron!er0 and they !aily deny that1 Since evidence means so little to them0 one
%onders %hy they don(t !o the %hole ho! and simply deny the %hole of plate tectonics too1
.erry Coyne(s Why Eol"tion is Tr"e offers a masterly treatment of the evidence from
!eo!raphical distriution 7as you(d expect from the senior author of the most authoritative recent
ook on speciation81 )e also hits the nail on the head %ith respect to the creationists( penchant for
i!norin! evidence %hen it doesn(t support the position that they know0 from Scripture0 has !ot to e
trueA (The io!eo!raphic evidence for evolution is no% so po%erful that # have never seen a
creationist ook0 article0 or lecture that has tried to refute it1 Creationists simply pretend that the
evidence doesn(t exist1( Creationists act as thou!h fossils provide the only evidence for evolution1
The fossil evidence is indeed very stron!1 Truckloads of fossils have een uncovered since 'ar%in(s
time0 and all this evidence either actively supports0 or is compatile %ith0 evolution1 "ore tellin!ly0
as # have already emphasiFed0 not a sin!le fossil contradicts evolution1 *evertheless0 stron! as the
fossil evidence is0 # a!ain %ant to emphasiFe that it is not the stron!est %e have1 Even if not a sin!le
fossil had ever een found0 the evidence from survivin! animals %ould still over%helmin!ly force
the conclusion that 'ar%in %as ri!ht1 The detective comin! on the scene of the crime after the event
can amass survivin! clues that are even more incontrovertile than fossils1 #n this chapter %e have
seen that the distriution of animals on islands and continents is exactly %hat %e should expect if
they are all cousins that have evolved from shared ancestors over very lon! periods1 #n the next
chapter %e shall compare modern animals %ith each other0 lookin! at the distriution of
characteristics in the animal kin!dom0 especially comparin! their seEuences of !enetic code0 and
shall come to the same conclusion1
LThe 1oya$e of the Bea$le1 +ictorian naturalists %ere !iven to value Hud!ements of this
kind in their ooks1 "y !randparents possessed a ird ook in %hich the entry on the cormorant
frankly e!an0 (There is nothin! to e said for this deplorale ird1(
L The rule seems to e that0 on islands0 i! animals !et smaller 7for example0 there %ere
d%arf elephants the hei!ht of a lar!e do! on "editerranean islands such as Sicily and Crete8 %hile
small animals !et i!!er 7as in the Galapa!os tortoises81 There are several theories for this diver!ent
tendency0 ut the details %ould take us too far afield1
L These para!raphs0 on !iant tortoises0 are extracted from an article that # %rote on a oat
called the Bea$le 7not the real one0 %hich is unfortunately lon! extinct8 in the Galapa!os
archipela!o0 and pulished in the G"ardian on 29 ,eruary 355=1
L -ike the modern (theory( of evolution0 it is an estalished fact in the normal sense of the
%ordA a theory in the first of the !ED(s definitions that # Euoted in Chapter 20 and renamed
L &las0 )olmes never said it 7Hust as Burns never %rote (for the sake of( &uld -an! Syne80
ut the allusion %orks ecause everyody thinks he did1
L #ndeed0 physical la%s of scalin! ensure that irds as i! as an elephant ird couldn(t
indul!e in po%ered0 flappin! fli!ht at all0 no matter ho% i! their %in! span1 This is ecause the
muscles needed to po%er such massive %in!s %ould need to e so i! they couldn(t lift their o%n
L #t is an arrestin! ima!eA South &merica and &frica speedin! a%ay from each other faster
than a man can s%im0 for forty days continuously1
C)&PTER 25
What a piece of %ork is the mammalian skeleton1 # don(t mean it is eautiful in itself0
althou!h # think it is1 # mean the fact that %e can talk aout (the( mammalian skeleton at allA the fact
that such a complicatedly interlockin! thin! is so !loriously different across the mammals0 in all its
parts0 %hile simultaneously ein! so oviously the same thin! throu!hout the mammals1 Our o%n
skeleton is familiar enou!h to need no picture0 ut look at this skeleton of a at1 #sn(t it fascinatin!
ho% every one has its o%n identifiale counterpart in the human skeletonB #dentifiale0 ecause of
the order in %hich they Hoin up to each other1 Only the proportions are different1 The at(s hands are
hu!ely enlar!ed 7relative to its total siFe0 of course8 ut noody could possily miss the
correspondence et%een our fin!ers and those lon! ones in the %in!s1 The human hand and the at
hand are oviously : no sane person could deny it : t%o versions of the same thin!1 The technical
term for this kind of sameness is (homolo!y(1 The at(s flyin! %in! and our !raspin! hand are
(homolo!ous(1 The hands of the shared ancestor : and the rest of the skeleton : %ere taken and
pulled0 or compressed0 part y part0 in different directions and y different amounts0 alon! different
descendant linea!es1
!at s&eleton
The same applies : althou!h %ith different proportions a!ain : in the %in! of a pterodactyl
7not a mammal0 ut the principle still holds0 %hich makes it all the more impressive81 This
pterodactyl(s %in! memrane is lar!ely orne y a sin!le fin!er0 %hat %e %ould call the (little(
fin!er or (pinky(1 # confess to a homolo!y:inspired neurosis aout so much %ei!ht ein! orne y
the fifth fin!er0 ecause in humans it seems so fra!ile1 Silly0 of course0 ecause to a pterodactyl the
fifth fin!er0 far from ein! (little(0 stretched most of the len!th of the ody0 and it presumaly %ould
have felt stout and stron!0 as our arm feels to us1 6et a!ain0 it !oes to illustrate the point # am
makin!1 The fifth fin!er is modified to ear the %in! memrane1 &ll the details have ecome
different0 ut it is still reco!niFaly the fifth fin!er ecause of its spatial relationship to the other
ones of the skeleton1 This lon!0 stout0 %in!:supportin! strut is (homolo!ous( to our little fin!er1
The %ord for (little fin!er( in pterodactylese means (ruddy !reat strut(1
#n addition to the true fliers : irds0 ats0 pterosaurs and insects : lots of other animals !lideA
a hait that mi!ht tell us somethin! aout the ori!ins of true fli!ht1 They have !lidin! memranes0
%hich need skeletal supportJ ut it doesn(t have to come from the fin!er ones as it does in the
%in!s of ats and pterosaurs1 ,lyin! sEuirrels 7t%o independent !roups of rodents80 and flyin!
phalan!ers 7&ustralian marsupials0 lookin! almost exactly like flyin! sEuirrels ut not closely
related8 stretch a memrane of skin et%een the arms and the le!s1 #ndividual fin!ers are not
reEuired to ear much load0 and they are not enlar!ed1 #0 %ith my little:fin!er neurosis0 %ould e
happier as a flyin! sEuirrel than as a pterodactyl0 ecause it feels (ri!ht( to use %hole arms and
%hole le!s to do load:earin! %ork1
7tero%atyl s&eleton
Overleaf is the skeleton of a so:called flyin! liFard0 another ele!ant forest !lider1 6ou can
immediately see that it is the ris0 rather than the fin!ers0 or the arms and le!s0 that have ecome
modified to ear the (%in!s( : the fli!ht memranes1 Once a!ain0 the resemlance of the skeleton as
a %hole to other verterate skeletons is completely clear1 6ou could !o throu!h every one0 one y
one0 identifyin!0 in each case0 the precise one to %hich it corresponds in the human or at or
pterosaur skeleton1
'Flyin' li-ar%' s&eleton
The colu!o0 or so:called (flyin! lemur(0 of the south:east &sian forests resemles the flyin!
sEuirrels and flyin! phalan!ers0 except that the tail0 as %ell as the arms and le!s0 is included in the
support structure of the fli!ht memrane1 That doesn(t feel ri!ht to me0 ecause # can(t ima!ine %hat
it is like to have a tail at all0 althou!h %e humans0 alon! %ith all the other (tail:less( apes0 have a
vesti!ial tail0 the coccyx0 uried eneath the skin1 &lmost tail:less as %e apes are0 it is hard for us to
ima!ine %hat it must e like to e a spider monkey0 %hose tail dominates the entire spinal column1
6ou can see from the picture on colour pa!e 3< ho% much lon!er it is even than the already lon!
arms and le!s1 &s in many *e% World monkeys 7indeed0 many *e% World mammals !enerally0
%hich is a curious fact0 hard to interpret80 the spider monkey(s tail is (prehensile(0 meanin! that it is
modified for !raspin!0 and it almost seems to end in an extra hand0 althou!h it is not homolo!ous to
a real hand0 and has no fin!ers1 #ndeed0 the spider monkey(s tail looks very much like an extra le! or
# proaly don(t need to spell out the messa!e a!ain1 The underlyin! skeleton is the same as
in the tail of any other mammal0 ut modified to do a different Ho1 Well0 the tail itself is not Euite
the sameA the spider monkey tail has an extra allo%ance of verterae0 ut the verterae themselves
are reco!niFaly the same kind of thin! as the verterae in any other tail0 includin! our o%n coccyx1
Can you ima!ine %hat it %ould e like to e a monkey %ith five !raspin! (hands( : one at the end of
each le! as %ell as at the end of each arm0 and a tail : from any of %hich you could happily han!B #
can(t1 But # kno% that the tail of a spider monkey is homolo!ous to my coccyx0 Hust as the
enormously lon! and stron! %in! one of a pterodactyl is homolo!ous to my little fin!er1
)ere(s another surprisin! fact1 & horse(s hoof is homolo!ous to the fin!ernail of your middle
fin!er 7or the toenail of your middle toe81 )orses %alk on tiptoe0 literally0 unlike us %hen %e %alk
on %hat %e call tiptoe1 They have almost entirely lost their other toes and fin!ers1 #n a horse0 the
homolo!ues of our index fin!er and our rin! fin!er0 and their hind:le! eEuivalents0 survive as tiny
(splint( ones0 Hoined to the (cannon( one0 and not visile outside the skin1 The cannon one is
homolo!ous to our middle metacarpal %hich is uried in our hand 7or metatarsal0 uried in our
foot81 The entire %ei!ht of the horse : very sustantial in the case of a Shire or a Clydesdale : is
orne on the middle fin!ers and middle toes1 The homolo!ies0 for example to our middle fin!ers or
those of a at0 are completely clear1 *oody could dout themJ and0 as if to ram the point home0
freak horses are sometimes orn %ith three toes on each le!0 the middle one servin! as a normal
(foot(0 the t%o side ones havin! miniature hooves 7see picture overleaf81
Can you see ho% eautiful it is0 this idea of near:indefinite modification over immensities of
time0 each modified form retainin! unmistakale traces of the ori!inalB # !lory in the litopterns0
extinct South &merican herivores0 not closely related to any modern animals0 and very different
from horses : except that they had almost identical le!s and hooves1 )orses 7in *orth &merica L8
and litopterns 7in South &merica0 %hich %as in those days a !i!antic island0 the Panama isthmus
ein! %ay in the future8 independently evolved exactly the same reduction of all the fin!ers and
toes except the middle ones0 and sprouted identical hooves on the ends of those1 Presumaly there
aren(t all that many %ays for a herivorous mammal to ecome a fast runner1 )orses and litopterns
hit upon the same %ay : reducin! all di!its except the middle one : and they carried it to the same
conclusion1 Co%s and antelopes hit upon another solutionA reducin! all ut t%o di!its1
7oly%atyli horse
The follo%in! statement sounds paradoxical ut you can see ho% it makes sense0 and also
ho% important it is as an oservation1 The skeletons of all mammals are identical0 ut their
individual ones are different1 The resolution of the paradox lies in my calculated usa!e of
(skeleton( for the assembla$e of ones0 in ordered attachment one to the other1 The shapes of
individual ones are not0 on this vie%0 properties of the (skeleton( at all1 (Skeleton(0 in this special
sense0 i!nores the shapes of individual ones0 and is concerned only %ith the order in %hich they
Hoin upA (one to his one( in the %ords of EFekiel0 and0 more vividly0 in the son! that is ased upon
the passa!eA
6our toe one connected to your foot one0
6our foot one connected to your ankle one0
6our ankle one connected to your le! one0
6our le! one connected to your knee one0
6our knee one connected to your thi!h one0
6our thi!h one connected to your hip one0
6our hip one connected to your ack one0
6our ack one connected to your shoulder one0
6our shoulder one connected to your neck one0
6our neck one connected to your head one0
# hear the %ord of the -ordK
The point is that this son! could apply to literally any mammal0 indeed any land verterate0
and in far more detail than these %ords su!!est1 ,or example your (head one(0 or skull0 contains
t%enty:ei!ht ones0 mostly Hoined to!ether in ri!id (sutures(0 ut %ith one maHor movin! one 7the
lo%er Ha%L81 &nd the %onderful thin! is that0 !ive or take the odd one here and there0 the same set
of t%enty:ei!ht ones0 %hich can clearly e laelled %ith the same names0 is found across all the
H("an s&(ll
Horse s&(ll
6our neck one connected to your occipital one
6our occiput connected to your parietal one
6our parietal connected to your frontal one
6our frontal one connected to your nasal one
1 1 1
6our 3@th one connected to your 3;th one 1 1 1
&ll this is the same0 re!ardless of the fact that the shapes of the particular ones are radically
different across the mammals1
What do %e conclude from all thisB We have here confined ourselves to modern animals0 so
%e are not seein! evolution in action1 We are the detectives0 come late to the scene1 &nd the pattern
of resemlances amon! the skeletons of modern animals is exactly the pattern %e should expect if
they are all descended from a common ancestor0 some of them more recently than others1 The
ancestral skeleton has een !radually modified do%n the a!es1 Some pairs of animals0 for example
!iraffes and okapis0 share a recent ancestor1 #t is not strictly correct to descrie a !iraffe as a
vertically stretched okapi0 for oth are modern animals1 But it %ould e a !ood !uess 7supported y
fossil evidence0 as it happens0 ut %e aren(t talkin! aout fossils in this chapter8 that the shared
ancestor proaly looked more like the okapi than the !iraffe1 Similarly0 impalas and !nus L are
close cousins of each other0 and sli!htly more distant cousins of !iraffes and okapis1 &ll four of
them are more distant cousins still of other cloven:hoofed animals0 such as pi!s and %artho!s
7%hich are cousins of each other and of peccaries81 &ll the cloven:hoofed animals are more distant
cousins of horses and Feras 7%hich don(t have cloven hooves and are close cousins of each other81
We can !o on as lon! as %e like0 racketin! pairs of cousins into !roups0 and !roups of !roups of
cousins0 and 7!roups of 7!roups of 7!roups of cousins8881 # have slipped into usin! rackets
automatically0 and you kno% Hust %hat they si!nify1 The meanin! of the rackets in the follo%in! is
immediately clear to you0 ecause you already kno% all aout cousins sharin! !randparents0 and
second cousins sharin! !reat:!randparents0 and so onA
7%olf fox87lion leopard87!iraffe okapi8 7impala !nu8
Everythin! points to a simple ranchin! tree of ancestry : a family tree1
# have implied that the tree of resemlances is really a family tree0 ut are %e forced to this
conclusionB &re there any alternative interpretationsB Well0 Hust arelyK The hierarchical pattern of
resemlances %as spotted y creationists in pre:'ar%inian times0 and they did have a non:
evolutionary explanation : an emarrassin!ly far:fetched one1 Patterns of resemlance0 in their
vie%0 reflected themes in the mind of the desi!ner1 )e had various ideas for ho% to make animals1
)is thou!hts ran alon! a mammal theme0 and0 independently0 they ran alon! an insect theme1
Within the mammal theme0 the desi!ner(s ideas %ere neatly and hierarchically isected into su:
themes 7say0 the cloven:hoofed theme8 and su:su:themes 7say0 the pi! theme81 There is a stron!
element of special pleadin! and %ishful thinkin! aout this0 and no%adays creationists seldom
resort to it1 #ndeed0 as %ith the evidence from !eo!raphical distriution0 %hich %e discussed in the
last chapter0 they rarely discuss comparative evidence at all0 preferrin! to stick to fossils0 %here they
have een tau!ht 7%ron!ly8 to think they are on promisin! !round1
To emphasiFe ho% odd the idea of a creator stickin! ri!idly to (themes( is0 reflect that any
sensile human desi!ner is Euite happy to orro% an idea from one of his inventions0 if it %ould
enefit another1 "aye there is a (theme( of aircraft desi!n0 %hich is separate from the (theme( of
train desi!n1 But a component of a plane0 say an improved desi!n for the readin! li!hts aove the
seats0 mi!ht as %ell e orro%ed for use in trains1 Why should it not0 if it serves the same purpose
in othB When motor cars %ere first invented0 the name (horseless carria!e( tells us %here some of
the inspiration came from1 But horse:dra%n vehicles don(t need steerin! %heels : you use reins to
steer horses : so the steerin! %heel had to have another source1 # don(t kno% %here it came from0
ut # suspect that it %as orro%ed from a completely different technolo!y0 that of the oat1 Before
ein! superseded y the steerin! %heel0 %hich %as introduced around the end of the nineteenth
century0 the ori!inal steerin! device of the car %as the tiller0 also orro%ed from oats0 ut moved
from the rear to the front of the vehicle1
#f feathers are a !ood idea %ithin the ird (theme(0 such that every sin!le ird0 %ithout
exception0 has them %hether it flies or not0 %hy do literally no mammals have themB Why %ould
the desi!ner not orro% that in!enious invention0 the feather0 for at least one atB The evolutionist(s
ans%er is clear1 &ll irds have inherited their feathers from their shared ancestor0 %hich had
feathers1 *o mammal is descended from that ancestor1 #t(s as simple as that1 L The tree of
resemlances is a family tree1 #t is the same kind of story for every ranch and every su:ranch
and every su:su:ranch of the tree of life1
*o% %e come to an interestin! point1 There are plenty of eautiful examples %here it looks0
superficially0 as thou!h ideas mi!ht have een (orro%ed( from one part of the tree and !rafted on to
another0 like an apple variety !rafted on to a stock1 & dolphin0 %hich is a small %hale0 looks
superficially like various kinds of lar!e fish1 One of these fish0 the dorado 7#oryphaena hipp"ris8 is
even sometimes called a (dolphin(1 'orados and true dolphins have the same streamlined shape0
suited to their similar %ays of life as fast hunters near the surface of the sea1 But their s%immin!
techniEue0 thou!h superficially similar0 %as not orro%ed from one y the other0 as you can Euickly
see if you look at the details1 &lthou!h oth derive their speed mostly from the tail0 the dorado0 like
all fish0 moves its tail from side to side1 But the true dolphin etrays its mammal history y eatin!
its tail up and do%n1 The side:to:side %ave travellin! do%n the ancestral fish ackone has een
inherited y liFards and snakes0 %hich could almost e said to (s%im( on land1 Contrast that %ith a
!allopin! horse or cheetah1 The speed comes from endin! of the spine0 as it does %ith fish and
snakesJ ut in mammals the spine ends up and do%n0 not side to side1 #t is an interestin! Euestion
ho% the transition %as made in the ancestry of mammals1 "aye there %as an intermediate sta!e0
%hich hardly ent its spine at all0 in either direction0 like a fro!1 On the other hand0 crocodiles are
capale of !allopin! 7fri!htenin!ly fast8 as %ell as usin! the liFard:like !ait more conventional
amon! reptiles1 The ancestors of mammals %ere nothin! like crocodiles0 ut maye crocodiles
sho% us ho% an intermediate ancestor mi!ht have comined the t%o !aits1
&ny%ay0 the ancestors of %hales and dolphins %ere fully paid:up land mammals0 %ho
surely !alloped across the prairies0 deserts or tundras %ith an up:and:do%n flexion of the spine1
&nd %hen they returned to the sea0 they retained their ancestral up:and:do%n spinal motion1 #f
snakes (s%im( on land0 dolphins (!allop( throu!h the seaK &ccordin!ly0 the fluke of a dolphin may
look superficially like the forked tail of a dorado0 ut it is set horiFontally0 %hereas the dorado(s tail
fins are ali!ned in the vertical plane1 There are numerous other respects in %hich the dolphin(s
history is %ritten all over it0 and # shall come to them in the chapter of that title1
There are other examples %here the superficial resemlance is so !reat that it seems Euite
hard to reHect the (orro%in!( hypothesis0 ut a closer inspection sho%s that %e must1 &nimals can
look so alike that you feel they must e related1 But it then turns out that the similarities0 thou!h
impressive0 are outnumered y the differences %hen you look at the %hole ody1 (Pill u!s( 7see
over8 are familiar little creatures0 %ith lots of le!s0 %ho haitually roll up into a protective all0 like
armadillos1 #ndeed0 this may e the ori!in of the -atin name Armadillidi"m4 That is the name of one
kind of (pill u!(0 %hich is a crustacean0 a %oodlouse0 related to shrimps ut livin! on land : %here
it etrays its recent aEuatic ancestry y reathin! %ith !ills0 %hich have to e kept moist1 But the
point of the story is that there is a completely different kind of (pill u!( %hich is not a crustacean at
all ut a millipede1 When you see them rolled up0 you(d think they %ere almost identical1 6et one is
a modified %oodlouse0 %hile the other is a modified 7modified in the same direction8 millipede1 #f
you unroll them and look carefully0 you %ill immediately see at least one important difference1 The
pill millipede has t%o pairs of le!s on most se!ments0 the pill %oodlouse only one1 #sn(t it eautiful0
all this endless modificationB & more detailed examination %ill sho% that0 in hundreds of respects0
the pill millipede really does resemle a more conventional millipede1 The resemlance to a
%oodlouse is superficial : conver!ent1
7ill "illi#e%e
7ill ,oo%lo(se
&lmost any Foolo!ist %ho %as not a specialist %ould say that the skull on the opposite pa!e
elon!s to a do!1 The specialist %ould discover that it isn(t actually a do! skull y notin! the t%o
prominent holes in the roof of the mouth1 These are tell:tale si!ns of marsupials0 the lar!e !roup of
mammals no%adays found mostly in &ustralia1 #t is in fact the skull of Thylacin"s0 the (Tasmanian
%olf(1 Thylacines and true do!s 7for example din!os0 %ith %hich they competed in &ustralia and
Tasmania8 have conver!ed on a very similar skull ecause they have 7had0 alas0 in the case of the
unfortunate thylacine8 a similar lifestyle1
# have already mentioned the ma!nificent marsupial mammal fauna of &ustralia0 in the
chapter on the !eo!raphical distriution of animals1 The relevant point for this chapter is the
repeated conver!ences et%een these marsupials and a !reat variety of opposite numers amon! the
(placental( 7i1e1 non:marsupial8 mammals0 %hich dominate the rest of the %orld1 Thou!h far from
identical0 even in superficial characteristics0 each marsupial in the illustration overleaf is sufficiently
similar to its placental eEuivalent : that is0 the placental that most closely practises the same (trade( :
to impress us0 ut certainly not sufficiently similar to su!!est (orro%in!( y a creator1
Thylaine '"ars(#ial ,ol$' or 'Tas"anian ti'er' s&(ll
The sexual shufflin! of the !enes in a !ene pool could e re!arded as a kind of orro%in! or
sharin! of !enetic (ideas(0 ut sexual recomination is confined %ithin one species and is therefore
irrelevant to this chapter0 %hich is aout comparisons et%een speciesA for example0 comparisons
et%een marsupial and placental mammals1 #nterestin!ly0 hi!h:level orro%in! of '*& is rife
amon! acteria1 #n a process that is sometimes re!arded as a kind of precursor to sexual
reproduction0 acteria : even Euite distantly related strains of acteria : s%ap '*& (ideas( %ith
promiscuous aandon1 (Borro%in! ideas( is indeed one of the main %ays y %hich acteria pick up
useful (tricks( such as resistance to particular antiiotics1
The phenomenon is often called y the rather unhelpful name of (transformation(1 That(s
ecause0 %hen it %as discovered in 293; y ,rederick Griffith0 noody understood aout '*&1
What Griffith found %as that a non:virulent strain of Streptococc"s could pick up virulence from a
completely different strain0 even thou!h that virulent strain %as dead1 *o%adays %e %ould say that
the non:virulent strain incorporated into its !enome some '*& from the dead virulent strain 7'*&
doesn(t care aout ein! (dead(0 it is Hust coded information81 #n the lan!ua!e of this chapter0 the
non:virulent strain (orro%ed( a !enetic (idea( from the virulent strain1 Of course0 acteria orro%in!
!enes from other acteria is a very different matter from a desi!ner orro%in! his o%n ideas from
one (theme( and re:usin! them in another theme1 *evertheless0 it is interestin! ecause0 if it %ere as
common in animals as it is in acteria0 it %ould make it harder to disprove the (desi!ner orro%in!(
hypothesis1 What if ats and irds ehaved like acteria in this respectB What if chunks of ird
!enome could e ferried across0 perhaps y acterial or viral infection0 and implanted in a at(s
!enomeB "aye a sin!le species of ats mi!ht suddenly sprout feathers0 the feather:codin! '*&
information havin! een orro%ed in a !enetic version of a computer(s (Copy and Paste(1
7laental an% "ars(#ial o##osite n("*ers
#n animals0 unlike acteria0 !ene transfer seems almost entirely confined to sexual con!ress
%ithin species1 #ndeed0 a species can pretty %ell e defined as a set of animals that en!a!e in !ene
transfer amon! themselves1 Once t%o populations of a species have een separated for lon! enou!h
that they can no lon!er exchan!e !enes sexually 7usually after an initial period of enforced
!eo!raphical separation0 as %e sa% in Chapter 980 %e no% define them as separate species0 and they
%ill never a!ain exchan!e !enes0 other than y the intervention of human !enetic en!ineers1 "y
collea!ue .onathan )od!kin0 Oxford(s Professor of Genetics0 kno%s of only three tentative
exceptions to the rule that !ene transfer is confined %ithin speciesA in nematode %orms0 in fruit
flies0 and 7in a i!!er %ay8 in delloid rotifers1
This last !roup is especially interestin! ecause0 uniEuely amon! maHor !roupin!s of
eucaryotes0 they have no sex1 Could it e that they have een ale to dispense %ith sex ecause they
have reverted to the ancient acterial %ay of exchan!in! !enesB Crossspecies !ene transfer seems to
e commoner in plants1 The parasitic plant dodder 7 #"sc"ta8 donates !enes to the host plants
around %hich it is ent%ined1 L
!%elloi% roti$er
# am undecided aout the politics of G" foods0 torn et%een the potential enefits to
a!riculture on the one hand and precautionary instincts on the other1 But one ar!ument # haven(t
heard efore is %orth a rief mention1 Today %e curse the %ay our predecessors introduced species
of animals into alien lands Hust for the fun of it1 The &merican !rey sEuirrel %as introduced to
Britain y a former 'uke of BedfordA a frivolous %him that %e no% see as disastrously
irresponsile1 #t is interestin! to %onder %hether taxonomists of the future may re!ret the %ay our
!eneration messed around %ith !enomesA transportin!0 for example0 (anti:freeFe( !enes from &rctic
fish into tomatoes to protect them from frost1 & !ene that !ives Hellyfish a fluorescent !lo% has een
orro%ed from them y scientists and inserted into the !enome of potatoes0 in the hope of makin!
them li!ht up %hen they need %aterin!1 # have even read of an (artist( %ho plans an (installation(
consistin! of luminous do!s0 !lo%in! %ith the aid of Hellyfish !enes1 Such deauchery of science in
the name of pretentious (art( offends all my sensiilities1 But could the dama!e !o furtherB Could
these frivolous caprices undermine the validity of future studies of evolutionary relationshipsB
&ctually # dout it0 ut perhaps the point is at least %orth raisin!0 in a precautionary spirit1 The
%hole point of the precautionary principle0 after all0 is to avoid future repercussions of choices and
actions that may not e oviously dan!erous no%1
# e!an the chapter %ith the verterate skeleton0 %hich is a lovely example of an invariant
pattern linkin! variale detail1 &lmost any other maHor !roup of animals %ould sho% the same kind
of thin!1 #(ll take Hust one other favourite exampleA the decapod crustaceans0 the !roup that includes
losters0 pra%ns0 cras and hermit cras 7%hich are not cras0 y the %ay81 The ody plan of all
crustaceans is the same1 Whereas our verterate skeleton consists of hard ones inside an other%ise
soft ody0 crustaceans have an exoskeleton consistin! of hard tues0 inside %hich the animal keeps
and protects its soft its1 The hard tues are Hointed and hin!ed0 in somethin! like the same %ay as
our ones are1 Think0 for example0 of the delicate hin!es in the le!s of a cra or loster0 and the
more roust hin!e of the cla%1 The muscles that po%er the pinch of a lar!e loster are inside the
tues that make up the cla%1 The eEuivalent muscles %hen a human hand pinches somethin! attach
to the ones that run throu!h the middle of the fin!er and thum1
-ike verterates0 ut unlike sea urchins or Hellyfish0 crustaceans are leftGri!ht symmetrical0
%ith a train of se!ments runnin! the len!th of the ody from head to tail1 The se!ments are the same
as each other in their underlyin! plan0 ut often differ in detail1 Each se!ment consists of a short
tue Hoined0 either ri!idly or y a hin!e0 to the t%o nei!hourin! se!ments1 &s %ith verterates0 the
or!ans and or!an systems of a crustacean sho% a repeat pattern as you move from front to rear1 ,or
example0 the main nerve trunk0 %hich runs the len!th of the ody on the ventral side 7not the dorsal
side0 as the verterate spinal cord does80 has a pair of !an!lia 7sort of mini:rains L8 in each
se!ment0 from %hich sprout nerves supplyin! the se!ment1 "ost of the se!ments have a lim on
each side0 each lim a!ain consistin! of a series of tues Hoined y hin!es1 Crustacean lims usually
terminate in a t%o:%ay ranch0 %hich in many cases you could call a cla%1 The head is se!mented
too althou!h0 as %ith the verterate head0 the se!mental pattern is more dis!uised here than in the
rest of the ody1 There are five pairs of lims lurkin! in the head0 althou!h it mi!ht sound a it
stran!e to call them lims since they are modified to ecome antennae or components of the Ha%
apparatus1 They are therefore usually called appenda!es rather than lims1 "ore or less invarialy0
the five se!mental appenda!es of the head0 readin! from the front0 consist of first antennae 7or
antennules80 second antennae 7often Hust called antennae80 mandiles0 first maxillae 7or maxillules8
and second maxillae1 The antennules and antennae are mostly en!a!ed in sensin! thin!s1 The
mandiles and maxillae are concerned %ith che%in!0 millin! or other%ise processin! food1 &s %e
proceed ack alon! the ody0 the se!mental appenda!es or lims are pretty variale0 the middle
ones often consistin! of %alkin! le!s0 %hile those sproutin! from the rearmost se!ments are often
pressed into service doin! other thin!s such as s%immin!1
#n a loster or a pra%n0 after the usual five head se!ment appenda!es0 the first ody se!ment
appenda!es are the cla%s1 The next four pairs are %alkin! le!s1 The se!ments earin! cla%s and
%alkin! le!s are unched to!ether as the thorax1 The rest of the ody is called the adomen1 #ts
se!ments0 at least until you reach the tip of the tail0 are the (s%immerets(0 feathery appenda!es that
help %ith s%immin!0 Euite importantly so in some delicately !raceful pra%ns1 #n cras0 the head
and thorax have mer!ed into a sin!le lar!e unit0 to %hich all the first ten pairs of lims are attached1
The adomen is douled ack under the headGthorax so that you can(t see it at all from aove1 But if
you turn a cra over0 you can clearly see the adomen(s se!mental pattern1 The picture elo% sho%s
the typical narro% adomen of a male cra1 The female adomen is %ider and resemles an apron0
%hich it is indeed called1 )ermit cras are unusual in that the adomen is asymmetrical 7to fit into
the empty mollusc shell %hich is its house80 and soft and unarmoured 7ecause the mollusc shell
provides protection81
4ale ra* sho,in' narro,, $ol%e%)*a& a*%o"en
To !et an idea of some of the %onderful %ays in %hich the crustacean ody is modified in
detail0 %hile the ody plan itself is not modified at all0 look at the set of dra%in!s opposite y the
famous nineteenth:century Foolo!ist Ernst )aeckel0 perhaps 'ar%in(s most devoted disciple in
Germany 7the devotion %as not reciprocated0 ut even 'ar%in %ould surely have admired
)aeckel(s drau!htsmanship81 .ust as %e did %ith the verterate skeleton0 look at each ody part of
these cras and crayfish0 and see ho%0 %ithout fail0 you can find its exact opposite numer in all the
rest1 Every it of the exoskeleton is Hoined to the (same( its0 ut the shapes of the its themselves
are very different1 Once a!ain0 the (skeleton( is invariant0 %hile its parts are anythin! ut1 &nd once
a!ain the ovious : # %ould say the only sensile : interpretation is that all these crustaceans have
inherited the plan of their skeleton from a common ancestor1 They have moulded the individual
components into a rich variety of shapes1 But the plan itself remains0 exactly as inherited from the
W)&T WO$-' '(&RC6 T)O"PSO* )&+E 'O*E W#T) & CO"P$TERB
#n 292@ the !reat Scottish Foolo!ist '(&rcy Thompson %rote a ook called !n Growth and
,orm0 in the last chapter of %hich he introduced his famous (method of transformations(1 L )e
%ould dra% an animal on !raph paper0 and then he %ould distort the !raph paper in a
mathematically specifiale %ay and sho% that the form of the ori!inal animal had turned into
another0 related animal1 6ou could think of the ori!inal !raph paper as a piece of ruer0 on %hich
you dra% your first animal1 Then the transformed !raph paper %ould e eEuivalent to the same
piece of ruer0 stretched or pulled out of shape in some mathematically defined %ay1 ,or example0
he took six species of cra and dre% one of them0 Geryon0 on ordinary !raph paper 7the undistorted
sheet of ruer81 )e then distorted his mathematical (ruer sheet( in five separate %ays0 to achieve
an approximate representation of the other five species of cra1 The details of the mathematics don(t
matter0 althou!h they are fascinatin!1 What you can clearly see is that it doesn(t take much to
transform one cra into another1 '(&rcy Thompson himself %asn(t very interested in evolution0 ut
it is easy for us to ima!ine %hat the !enetic mutations %ould have to do in order to rin! aout
chan!es like this1 That doesn(t mean %e should think of Geryon0 or any other one of these six cras0
as ein! ancestral to the others1 *one of them %as0 and in any case that is not the point1 The point is
that %hatever the ancestral cra looked like0 transformations of this kind could chan!e any one of
these six species 7or a putative ancestor8 into any other1
Hae&el's r(staeans. 5rnst Hae&el ,as a %istin'(ishe% >er"an -oolo'ist an% an
e/ellent -oolo'ial artist.
Evolution never happened y takin! one adult form and coaxin! it into the shape of another1
Rememer that every adult !ro%s as an emryo1 The mutations selected %ould have %orked in the
developin! emryo y chan!in! the rate of !ro%th of parts of the ody relative to other parts1 #n
Chapter @ %e interpreted the evolution of the human skull as a series of chan!es in the rates of
!ro%th of some parts relative to other parts0 controlled y !enes in the developin! emryo1 We
should expect0 therefore0 that if %e dra% a human skull on a sheet of (mathematical ruer(0 it
should e possile to distort the ruer in some mathematically tidy %ay and achieve an
approximate likeness to the skull of a close cousin0 such as a chimpanFee0 or : perhaps %ith a i!!er
distortion : a more distant cousin such as a aoon1 &nd this is Hust %hat '(&rcy Thompson
sho%ed1 *ote0 once a!ain0 that it %as an aritrary decision to dra% the human skull first0 and then
transform it into the chimpanFee and the aoon1 )e could eEually %ell have dra%n0 say0 the
chimpanFee first and then %orked out the necessary distortions to make the human and the aoon1
Or0 more interestin!ly for a ook on evolution0 %hich his %as not0 he mi!ht have dra%n0 say0 an
A"stralopithec"s skull first on the undistorted ruer0 and %orked out ho% to transform it to make a
modern human skull1 This %ould surely have %orked Hust as %ell as the pictures aove0 and it
%ould have een evolutionarily meanin!ful in a more direct %ay1
D'Ary Tho"#son's ra* 'trans$or"ations'
D'Ary Tho"#son's s&(ll 'trans$or"ation'
&t the e!innin! of this chapter # introduced the idea of (homolo!y(0 usin! the arms of ats
and humans as an example1 #ndul!in! an idiosyncratic use of lan!ua!e0 # said that the skeletons
%ere identical %hile the ones %ere different1 '(&rcy Thompson(s transformations furnish us %ith a
%ay to make this idea more precise1 #n this formulation0 t%o or!ans : for example0 at hand and
human hand : are homolo!ous if it is possile to dra% one on a sheet of ruer and then distort the
ruer to make the other one1 "athematicians have a %ord for thisA (homeomorphic(1L
Ioolo!ists reco!niFed homolo!y in pre:'ar%inian times0 and pre:evolutionists %ould
descrie0 say0 at %in!s and human hands as homolo!ous1 #f they had kno%n enou!h mathematics0
they %ould have een happy to use the %ord (homeomorphic(1 #n post:'ar%inian times0 %hen it
ecame !enerally accepted that ats and humans share a common ancestor0 Foolo!ists started to
define homolo!y in evolutionary terms1 )omolo!ous resemlances are those inherited from the
shared ancestor1 The %ord (analo!ous( came to e used for resemlances due to shared function0 not
ancestry1 ,or example0 a at %in! and an insect %in! %ould e descried as analo!ous0 as opposed
to the homolo!ous at %in! and human arm1 #f %e %ant to use homolo!y as evidence for the fact of
evolution0 %e can(t use evolution to define it1 ,or this purpose0 therefore0 it is convenient to revert to
the pre:evolutionary definition of homolo!y1 The at %in! and human arm are homeomorphicA you
can transform one into the other y distortin! the ruer on %hich it is dra%n1 6ou cannot
transform a at %in! into an insect %in! in this %ay0 ecause there are no correspondin! parts1 The
%idespread existence of homeomorphisms0 %hich are not defined in terms of evolution0 can e used
as evidence for evolution1 #t is easy to see ho% evolution could !o to %ork on any verterate arm
and transform it into any other verterate arm0 simply y chan!in! relative rates of !ro%th in the
Ever since ecomin! acEuainted %ith computers as a !raduate student in the 29<5s0 # have
%ondered %hat '(&rcy Thompson mi!ht have done %ith a computer1 The Euestion ecame pressin!
in the 29;5s0 %hen affordale computers %ith screens 7as opposed to Hust paper printers8 ecame
common1 'ra%in! on stretched ruer and then distortin! the dra%in! surface in a mathematical
%ay : it %as Hust be$$in$ for the computer treatmentK # su!!ested that Oxford $niversity should id
for a !rant to employ a pro!rammer to put '(&rcy Thompson(s transformations on a computer
screen0 and make them availale in a user:friendly manner1 We !ot the money0 and employed Will
&tkinson0 a first:class pro!rammer and iolo!ist0 %ho ecame a friend and an adviser to me on my
o%n pro!rammin! proHects1 Once he had solved the difficult prolem of pro!rammin! a rich
repertoire of mathematical distortions of the (ruer(0 it %as then a relatively simple task for him to
incorporate this mathematical %iFardry into a iomorph:style artificial selection pro!ram0 similar to
my o%n (iomorph( pro!rams0 here descried in Chapter 31 &s %ith my pro!rams0 the (player( %as
confronted %ith a screen full of animal forms0 and invited to choose one of them for (reedin!(0
!eneration after !eneration1 Once a!ain there %ere (!enes( that persisted throu!h the !enerations0
and once a!ain the !enes influenced the form of the (animals(1 But in this case0 the %ay the !enes
influenced animal form %as y controllin! the distortion of the (ruer( on %hich an animal(s form
had een dra%n1 Theoretically0 therefore0 it should have een possile to start %ith0 say0 an
A"stralopithec"s skull dra%n on the undistorted (ruer(0 and reed your %ay throu!h creatures %ith
pro!ressively lar!er raincases and pro!ressively shorter muFFles : pro!ressively more human:like0
in other %ords1 #n practice it proved very difficult to do anythin! like that0 and # think the fact is0 in
itself0 interestin!1
# think one reason it %as difficult is0 yet a!ain0 that '(&rcy Thompson(s transformations
chan!e one ad"lt form into another adult form1 &s # emphasiFed in Chapter ;0 that is not ho% !enes
in evolution %ork1 Every individual animal has a developmental history1 #t starts as an emryo and
!ro%s0 y disproportionate !ro%th of different parts of the ody0 into an adult1 Evolution is not a
!enetically controlled distortion of one adult form into anotherJ it is a !enetically controlled
alteration in a developmental pro!ram1 .ulian )uxley 7!randson of T1)1 and rother of &ldous8
reco!niFed this %hen0 soon after pulication of the first edition of '(&rcy Thompson(s ook0 he
modified the (method of transformations( to study the %ay early emryos turn into later emryos or
adults1 That(s all # %ant to say aout '(&rcy Thompson(s method of transformations here1 #(ll return
to the topic in the final chapter0 to make a related point1
Comparative evidence has al%ays0 as # su!!ested at the e!innin! of this chapter0 told even
more compellin!ly than fossil evidence in favour of the fact of evolution1 'ar%in himself took a
similar vie%0 at the end of his chapter in !n the !ri$in of Species on the ("utual &ffinities of
Or!anic Bein!s(A
,inally0 the several classes of facts %hich have een considered in this chapter0 seem to me
to proclaim so plainly0 that the innumerale species0 !enera0 and families of or!anic ein!s0 %ith
%hich this %orld is peopled0 have all descended0 each %ithin its o%n class or !roup0 from common
parents0 and have all een modified in the course of descent0 that # should %ithout hesitation adopt
this vie% even if it %ere unsupported y other facts or ar!uments1
What 'ar%in didn(t : couldn(t : kno% is that the comparative evidence ecomes even more
convincin! %hen %e include molecular !enetics0 in addition to the anatomical comparisons that
%ere availale to him1
.ust as the verterate skeleton is invariant across all verterates %hile the individual ones
differ0 and Hust as the crustacean exoskeleton is invariant across all crustaceans %hile the individual
(tues( vary0 so the '*& code is invariant across all livin! creatures0 %hile the individual !enes
themselves vary1 This is a truly astoundin! fact0 %hich sho%s more clearly than anythin! else that
all livin! creatures are descended from a sin!le ancestor1 *ot Hust the !enetic code itself0 ut the
%hole !eneGprotein system for runnin! life0 %hich %e dealt %ith in Chapter ;0 is the same in all
animals0 plants0 fun!i0 acteria0 archaea and viruses1 What varies is %hat is %ritten in the code0 not
the code itself1 &nd %hen %e look comparatively at %hat is %ritten in the code : the actual !enetic
seEuences in all these different creatures : %e find the same kind of hierarchical tree of
resemlance1 We find the same familytree : aleit much more thorou!hly and convincin!ly laid out
: as %e did %ith the verterate skeleton0 the crustacean skeleton0 and indeed the %hole pattern of
anatomical resemlances throu!h all the livin! kin!doms1
#f %e %ant to %ork out ho% closely related any pair of species is : say0 ho% close a
hed!eho! is to a monkey : the ideal %ould e to look at the complete molecular texts of every !ene
of oth species0 and compare every Hot and tittle0 as a ilical scholar mi!ht compare t%o scrolls or
fra!ments of #saiah1 But it is time:consumin! and expensive1 The )uman Genome ProHect took
aout ten years0 representin! many person:centuries1 &lthou!h it %ould no% e possile to achieve
the same result in a fraction of the time0 it %ould still e a lar!e and expensive undertakin!0 as
%ould the hed!eho! !enome proHect1 -ike the &pollo moon landin!s0 and like the -ar!e )adron
Collider 7%hich has Hust een s%itched on in Geneva as # %rite : the !i!antic scale of this
international endeavour moved me to tears %hen # visited80 the complete decipherin! of the human
!enome is one of those achievements that makes me proud to e human1 # am deli!hted that the
chimpanFee !enome proHect has no% een successfully accomplished0 and the eEuivalent for
various other species1 #f the present rate of pro!ress continues 7see ()od!kin(s -a%( elo%80 it %ill
soon e economically feasile to seEuence the !enome of every pair of species %hose closeness of
cousinship %e mi!ht %ant to measure1 "ean%hile0 for the most part %e have to resort to samplin!
particular parts of their !enomes0 and it %orks pretty %ell1
We can sample y pickin! out a fe% choice !enes 7or proteins0 %hose seEuences are directly
translated from !enes8 and comparin! them across species1 #(ll come to that in a moment1 But there
are other %ays of doin! a kind of crude0 automatic samplin!0 and the technolo!ies to do that have
een around for lon!er1 &n early method0 %hich %orks surprisin!ly %ell0 exploits the immune
system of raits 7you could actually use any animal you like0 ut raits do the Ho nicely81 &s part
of the ody(s natural defence a!ainst patho!ens0 the rait(s immune system manufactures
antiodies a!ainst any forei!n protein that enters the loodstream1 .ust as you could tell that # have
had %hoopin! cou!h y lookin! at the antiodies in my lood0 so you can tell %hat a rait has
een exposed to in the past y lookin! at its immune response in the present1 The antiodies present
in the rait constitute a history of the natural shocks to %hich its flesh has een heir : includin!
artificially inHected proteins1 #f you inHect0 say0 a chimpanFee protein into a rait0 the antiodies
that it makes %ill suseEuently attack the same protein if it is inHected a!ain1 But suppose your
second inHection is of the eEuivalent protein0 not from a chimpanFee ut from a !orillaB The rait(s
prior exposure to the chimpanFee protein %ill have partially forearmed it a!ainst the !orilla version0
ut the reaction %ill e %eaker1 &nd it %ill also have forearmed it a!ainst the kan!aroo version of
the protein0 ut the reaction %ill e %eaker still0 !iven that the kan!aroo is much less closely related
to the chimpanFee that did the primin! than the !orilla is1 The stren!th of the rait(s immune
response to a suseEuent inHection of a protein is a measure of the resemlance of that protein to the
ori!inal to %hich the rait %as first exposed1 #t %as y this method0 usin! raits0 that +incent
Sarich and &llan Wilson0 at the $niversity of California at Berkeley0 demonstrated in the 29<5s that
humans and chimpanFees are much more closely related to each other than anyody had previously
There are also methods that use the !enes themselves0 comparin! them across species
directly rather than comparin! the proteins they encode1 One of the oldest and most effective of
these methods is called '*& hyridiFation1 '*& hyridiFation is usually %hat lies ehind those
statements one often sees alon! the lines ofA ()umans and chimpanFees share 9; per cent of their
!enes1( There is some confusion0 y the %ay0 aout exactly %hat is meant y percenta!e fi!ures
such as these1 *inety:ei!ht per cent of what is identicalB The exact fi!ure depends on ho% lar!e the
units are that %e are countin!1 & simple analo!y makes this clear0 and it does so in an interestin!
%ay0 ecause the differences et%een the analo!y and the real thin! are as revealin! as the
similarities1 Suppose %e have t%o versions of the same ook and %e %ant to compare them1
Perhaps it is the ook of 'aniel0 and %e %ant to compare the canonical version %ith an ancient
scroll that has Hust een discovered in a cave overlookin! the 'ead Sea1 What percenta!e of the
chapters of the t%o ooks are identicalB Proaly Fero0 for it takes only one discrepancy0 any%here
in the %hole chapter0 for us to say the t%o are not identical1 What percenta!e of their sentences are
identicalB The percenta!e %ill no% e much hi!her1 Even hi!her %ill e the percenta!e of %ords
that are identical0 ecause %ords have fe%er letters than sentences : fe%er opportunities to ust the
identity1 But a %ord resemlance is still roken if any one letter in the %ord differs1 Therefore0 if
you line the t%o texts up side y side and compare them letter y letter0 the percenta!e of identical
letters %ill e even hi!her than the percenta!e of identical %ords1 So an estimate like (9; per cent in
common( doesn(t mean anythin! unless %e specify the siFe of the unit %e are comparin!1 &re %e
countin! chapters0 %ords0 letters or %hatB &nd the same is true %hen %e compare '*& from t%o
species1 #f you are comparin! %hole chromosomes0 the percenta!e shared is Fero0 ecause it only
takes one tiny difference0 some%here alon! the chromosomes0 to define the chromosomes as
The often:Euoted fi!ure of aout 9; per cent for the shared !enetic material of humans and
chimps actually refers neither to numers of chromosomes nor to numers of %hole !enes0 ut to
numers of '*& (letters( 7technically0 ase pairs8 that match each other %ithin the respective human
and chimp !enes1 But there is a pitfall1 #f you do the linin! up naively0 a missin$ letter 7or an added
letter80 as opposed to a mistaken letter0 %ill cause all suseEuent letters to mismatch0 ecause they
%ill then all e sta!!ered0 one step out 7until there is a mistake in the other direction to rin! them
ack into step a!ain81 #t is clearly unfair to let the estimate of discrepancies e inflated in this %ay1
& scholar(s eye0 scannin! t%o scrolls of 'aniel0 automatically copes %ith this0 in a %ay that is hard
to Euantify1 )o% can %e do it %ith '*&B This is %here %e leave our analo!y %ith ooks and
scrolls and !o strai!ht to the real thin! ecause0 as it happens0 the real thin! : '*& : is easier to
understand than the analo!yK
#f you !radually heat '*&0 there comes a point : some%here around ;=de!C : %hen the
ondin! et%een the t%o strands of the doule helix reaks0 and the t%o helices separate1 6ou can
think of ;=de!C0 or %hatever the temperature turns out to e0 as a (meltin! point(1 #f you let it cool
a!ain0 each sin!le helix spontaneously Hoins up a!ain %ith another sin!le helix0 or fra!ment of
sin!le helix0 %herever it finds one %ith %hich it can pair0 usin! the ordinary ase:pairin! rules of
the doule helix1 6ou mi!ht think that this %ould al%ays e the partner from %hich it lately
separated0 and %ith %hich0 of course0 it is perfectly matched1 #ndeed it could e0 ut it usually isn(t
as tidy as that1 ,ra!ments of '*& %ill find other fra!ments %ith %hich they can pair0 and they %ill
usually not e exactly their ori!inal partners1 &nd indeed0 if you add separated fra!ments of '*&
from another species0 fra!ments of the sin!le strands are Euite capale of Hoinin! up %ith fra!ments
of sin!le strands from the %ron! species0 in Hust the same %ay as they %ill Hoin up %ith sin!le
strands from the ri!ht species1 Why should they notB #t is the remarkale conclusion of the Watson:
Crick molecular iolo!y revolution that '*& is Hust '*&1 #t doesn(t (care( %hether it is human
'*&0 chimp '*& or apple '*&1 ,ra!ments %ill happily pair off %ith complementary fra!ments
%herever they find them1 *evertheless0 the stren!th of ondin! is not al%ays eEual1 Sin!le:stranded
len!ths of '*& ond more ti!htly %ith matchin! sin!le strands than they do %ith less similar
sin!le strands1 This is ecause more of the (letters( of the '*& 7Watson and Crick(s (ases(8 find
themselves opposite partners %ith %hich they cannot pair1 The ondin! of the strands is therefore
%eakened : like a Fip fastener %ith some teeth missin!1
)o% shall %e measure this stren!th of ondin!0 after fra!ments from different species have
found each other and unitedB By an almost ludicrously simple method1 We measure the (meltin!
point( of the onds1 6ou rememer # said that the meltin! point of doule:stranded '*& is aout
;=de!C1 This is true of normal0 properly matched doule:stranded '*&0 as %hen a strand of human
'*& is (melted( a%ay from a complementary strand of human '*&1 But %hen the ondin! is
%eaker : as %hen a human strand has onded %ith a chimpanFee strand : a sli!htly lo%er
temperature is sufficient to reak the ond1 &nd %hen human '*& has onded %ith '*& from a
more distant cousin like a fish or a toad0 an even lo%er temperature suffices to separate them1 The
difference et%een the meltin! point %hen a strand is onded to one of its o%n kind0 and the
meltin! point %hen it is onded to a strand from another species0 is our measure of the !enetic
distance et%een the t%o species1 &s a rule of thum0 each decrease y 2de! Celsius in the (meltin!
point( is approximately eEuivalent to a drop of 2 per cent in the numer of '*& letters matched 7or
an increase of 2 per cent in the numer of missin! teeth in the Fip fastener81
There are complications in the method0 %hich # haven(t !one into0 and tricky prolems0
%hich have in!enious solutions1 ,or instance0 if you mix human %ith chimp '*&0 much of the
fra!mented human '*& %ill ond %ith other human '*& fra!ments0 and much of the chimp '*&
%ill ond %ith its o%n kind1 )o% do you separate off the hyrid '*&0 %hose (meltin! point( is
%hat you really %ant to measure0 from the (same:kind( '*&B The ans%er is y a clever trick
involvin! previous radioactive laellin!1 But the details %ould take us too far off our path1 The
main point here is that '*& hyridiFation is the techniEue that leads scientists to fi!ures like 9; per
cent for the !enetic similarity et%een humans and chimpanFees0 and it yields predictaly lo%er
percenta!es as you move to more distantly related pairs of animals1
The ne%est method of measurin! the similarity et%een a pair of matchin! !enes from
different species is the most direct0 and the most expensiveA actually read the seEuence of letters in
the !enes themselves0 usin! the same methods as %ere used for the )uman Genome ProHect1
&lthou!h it is still expensive to compare the entire !enome0 you can !et a !ood approximation y
comparin! a sample of !enes0 and this is no% increasin!ly done1
Whichever techniEue %e use for measurin! similarity et%een t%o species0 %hether it is
rait antiodies0 or meltin! points0 or direct seEuencin!0 the next step is pretty much the same1
)avin! otained a sin!le numer representin! the similarity et%een each pair of species0 %e then
place the fi!ures in a tale1 Take a set of species and %rite their names0 in the same order0 as oth
the column headin!s and the ro% headin!s1 Then place the percenta!e similarities in the appropriate
cells1 The tale %ill e trian!ular 7half of a sEuare8 ecause0 for example0 the percenta!e similarity
et%een human and do! %ill e the same as the similarity et%een do! and human1 So if you filled
in all of a sEuare tale each of the t%o halves either side of the dia!onal %ould mirror the other1
*o%0 %hat sort of results should %e expectB On the evolution model %e should predict that
you(ll find yourself puttin! a hi!h score in the cell connectin! human and chimpanFeeJ a lo%er
score in the cell connectin! human and do!1 The humanGdo! cell should theoretically have an
identical resemlance score to the chimpanFeeG do! cell ecause humans and chimpanFees have
exactly the same de!ree of relation to do!s1 #t should e identical0 too0 to the monkeyG do! cell and
the lemurGdo! cell1 This is ecause humans0 chimpanFees0 monkeys and lemurs are all connected to
the do! via their common ancestor0 an early primate 7%hich proaly looked a it like a lemur81 The
same score should sho% up in the humanGcat0 chimpanFeeGcat0 monkeyGcat and lemurGcat cells0
ecause cats and do!s are related to all primates via the shared ancestor of all carnivores1 There
should e a much lo%er score : ideally eEually lo% : in all the cells unitin!0 say0 a sEuid %ith any
mammal1 &nd it shouldn(t matter %hich mammal you choose0 since all are eEually distant from a
These are stron! theoretical expectations0 ut there is no reason %hy0 in practice0 they
should not e violated1 #f they %ere violated0 it %ould e evidence a!ainst evolution1 What actually
happens turns out to e : %ithin statistical mar!ins of error : Hust %hat %e should expect on the
assumption that evolution has happened1 This is another %ay of sayin! that0 if you put the !enetic
distances et%een pairs of species on the lims of a tree0 everythin! adds up in a satisfyin! %ay1 Of
course the addin! up is not Euite perfect1 *umerical expectations in iolo!y are seldom realiFed
%ith etter than approximate accuracy1
Comparative '*& 7or protein8 evidence can e used to decide : on the evolutionary
assumption : %hich pairs of animals are closer cousins than %hich others1 What turns this into
extremely po%erful evidence for evolution is that you can construct a tree of !enetic resemlances
separately for each !ene in turn1 &nd the important result is that every !ene delivers approximately
the same tree of life1 Once a!ain0 this is exactly %hat you %ould expect if you %ere dealin! %ith a
true family tree1 #t is not %hat you %ould expect if a desi!ner had surveyed the %hole animal
kin!dom and picked and chosen : or (orro%ed( : the est proteins for the Ho0 %herever in the
animal kin!dom they mi!ht e found1
The earliest lar!e:scale study alon! these lines %as done y a !roup of !eneticists in *e%
Iealand led y Professor 'avid Penny1 Penny(s !roup took five !enes %hich0 althou!h not identical
across all mammals0 %ere similar enou!h to have earned the same name in all1 The details don(t
matter0 ut for the record the five !enes %ere those for haemo!loin &0 haemo!loin B
7haemo!loins !ive lood its red colour80 firinopeptide &0 firinopeptide B 7firinopeptides are
used in clottin! lood8 and cytochrome C 7%hich plays an important role in cellular iochemistry81
They chose eleven mammals to compareA rhesus monkey0 sheep0 horse0 kan!aroo0 rat0 rait0 do!0
pi!0 human0 co% and chimpanFee1
Penny and his collea!ues thou!ht statistically1 They %anted to calculate the proaility that0
purely y chance0 t%o molecules %ould yield the same family tree0 if evolution %asn(t true1 So they
tried to ima!ine all possile trees that could terminate in eleven descendants1 #t(s a surprisin!ly lar!e
numer1 Even if you limit yourself to (inary trees( 7that is0 trees %ith ranches that only ifurcate :
no tri:furcatin! or hi!her:furcatin!80 the total numer of possile trees is more than 4> million1 The
scientists patiently looked at every one of the 4> million trees and compared each one %ith the other
4409990999 trees1 *o0 of course they didn(tK #t %ould take too much computer time1 They did0
ho%ever0 devise a clever statistical approximation0 a shortcut eEuivalent to that mammoth
This is ho% the method of approximation %orked1 They took the first of the five !enes0 say
haemo!loin:& 7in all cases # use the name of the protein to stand for the !ene that codes for that
protein81 Of all those millions of trees0 they %anted to find %hich %as the most (parsimonious(
%here haemo!loin:& %as concerned1 Parsimonious here means (needin! to postulate the minimum
amount of evolutionary chan!e(1 ,or example0 all those thousands of trees that assumed that the
closest cousin to a human %as a kan!aroo %hile humans and chimpanFees are more distantly
related0 proved to e very unparsimonious treesA they needed to assume a lot of evolutionary
chan!e0 in order to yield the result that kan!aroos and humans had a recent common ancestor1
)aemo!loin:&(s verdict %ould e alon! these linesA
This is a terrily unparsimonious tree1 *ot only do # have to put in lots of mutational %ork
in order to end up so different in humans and kan!aroos0 despite our close cousinship accordin! to
this tree0 # also have to put in lots of mutational %ork in the other direction0 in order to ensure that0
despite their !reat separation on this particular tree0 humans and chimps someho% ended up %ith
such similar haemo!loin:&1 # vote a!ainst this tree1
)aemo!loin:& delivers a verdict of this kind0 some verdicts more favourale than others0
on each of the 4> million trees0 and finally ends up choosin! a fe% doFen top:rankin! trees1 Of each
of these top:rankin! trees0 haemo!loin:& %ould say somethin! like thisA
This tree puts humans and chimpanFees as close cousins0 and it puts sheep and co%s as close
cousins0 and it puts kan!aroos out on a lim1 This turns out to e a very !ood tree0 ecause it makes
me do hardly any mutational %ork at all to explain the evolutionary chan!es1 This is an excellently
parsimonious tree1 #t !ets the haemo!loin:& voteK
Of course0 it %ould have een nice if haemo!loin:&0 and every other !ene0 could have
come up %ith a sin!le most parsimonious tree0 ut that is too much to ask1 &mon! the 4> million
trees0 it is only to e expected that several sli!htly different trees should tie for haemo!loin:&(s
top:rankin! slot1
*o%0 ho% aout haemo!loin:BB )o% aout cytochrome:CB Each one of the five proteins
is entitled to its o%n separate vote0 to find its o%n preferred 7that is0 most parsimonious8 trees from
amon! the 4> million trees1 #t %ould e perfectly possile for cytochrome:C to come up %ith a
completely different vote on %hich is the most parsimonious tree1 #t could turn out that the
cytochrome:C of humans really is very similar to that of kan!aroos0 and very different from that of
chimpanFees1 ,ar from salutin! the close pairin! of sheep and co% discerned y haemo!loin:&0
cytochrome:C mi!ht find that it hardly needs to mutate at all in order to place sheep very close to0
say0 monkeys0 and in order to place co%s very close to raits1 On the creation hypothesis there is
no reason %hy that shouldn(t happen1 But %hat Penny and his collea!ues actually found %as that
there %as astonishin!ly hi!h a!reement amon! all five proteins 7and they used yet more clever
statistics to sho% ho% unlikely such concordance %ould e y chance81 &ll five proteins (voted( for
pretty much the same suset of trees from amon! the 4> million possile trees1 This is0 of course0
exactly %hat %e %ould expect on the assumption that there really is only one true tree relatin! all
eleven animals0 and it is the family treeA the tree of evolutionary relationships1 What is more0 the
consensus tree that the five molecules all voted for turned out to e the same as Foolo!ists had
already %orked out on anatomic and palaeontolo!ical0 not molecular0 !rounds1
The Penny study %as pulished in 29;30 Euite a %hile a!o no%1 The intervenin! years have
seen a prolific multiplication of detailed evidence on the exact seEuences of !enes of lots and lots of
species of animals and plants1 &!reement on the most parsimonious trees no% extends far eyond
the eleven species and five molecules that Penny and his collea!ues studied1 Theirs %as Hust a nice
example0 over%helmin! as their statistical evidence proved1 The sum total of !enetic seEuence data
no% availale puts the matter eyond all conceivale dout1 ,ar more convincin!ly even than the
7also hi!hly convincin!8 fossil evidence0 the evidence from comparisons amon! !enes is
conver!in!0 rapidly and decisively0 on a sin!le !reat tree of life1 &ove is a tree for the eleven
species of the Penny study0 %hich represents a modern consensus vote from many different parts of
the mammalian !enome1 #t is the consistency of a!reement amon! all the different !enes in the
!enome that !ives us confidence0 not only in the historical accuracy of the consensus tree itself0 ut
also in the fact that evolution has occurred1
Fa"ily tree $or 7enny's ele.en s#eies
#f molecular !enetic technolo!y continues to expand at its present exponential rate0 y the
year 35=5 derivin! the complete seEuence of an animal(s !enome %ill e cheap and Euick0 scarcely
any more troule than takin! its temperature or its lood pressure1 Why do # say that !enetic
technolo!y is expandin! exponentiallyB Could %e even measure itB There is a parallel in computer
technolo!y called "oore(s -a%1 *amed after Gordon "oore0 one of the founders of the #ntel
computer chip company0 it can e expressed in various %ays ecause several measures of computer
po%er are linked to each other1 One version of the la% states that the numer of units that can e
packed into an inte!rated circuit of a !iven siFe doules every ei!hteen months to t%o years or so1 #t
is an empirical la%0 meanin! that0 rather than derivin! from some piece of theory0 it Hust turns out to
e true %hen you measure the data1 #t has held !ood over a period of aout fifty years so far0 and
many experts think it %ill do so for at least a fe% more decades1 Other exponential trends0 %ith a
similar doulin! time0 %hich can e re!arded as versions of "oore(s -a%0 include the increase in
speed of computation0 and siFe of memory0 per unit cost1 Exponential trends al%ays lead to startlin!
results0 as 'ar%in demonstrated %hen0 %ith the aid of his mathematician son Geor!e0 he took the
elephant as an example of a slo%:reedin! animal and sho%ed that0 in Hust a fe% centuries of
unrestricted exponential !ro%th0 the descendants of Hust one pair of elephants %ould carpet the
earth1 *eedless to say0 population !ro%th of elephants is not0 in practice0 exponential1 #t is limited
y competition for food and space0 y disease0 and y many other thin!s1 That0 indeed0 %as
'ar%in(s %hole point0 for that is %here natural selection steps in1
But "oore(s -a% really has remained in force0 at least approximately0 for fifty years1
&lthou!h noody has a very clear idea %hy0 various measures of computer po%er actually have
increased exponentially in practice0 %here 'ar%in(s elephant trend is exponential only in theory1 #t
occurred to me that there mi!ht e a similar la% in force for !enetic technolo!y and the seEuencin!
of '*&1 # su!!ested it to .onathan )od!kin0 Oxford(s Professor of Genetics 7%ho had once een an
under!raduate pupil of mine81 To my deli!ht0 it turned out that he had already thou!ht of it : and
measured it0 in preparation for a lecture at his old school1 )e estimated the cost of seEuencin! a
standard len!th of '*& at four dates in history0 29<=0 29@=0 299= and 35551 # inverted his fi!ures
to (an!s for the uck(0 or ()o% much '*& could you seEuence for PS20555B( # plotted the fi!ures
on a lo!arithmic scale0 chosen ecause an exponential trend %ill al%ays sho% up as a strai!ht line
%hen plotted lo!arithmically1 Sure enou!h0 )od!kin(s four points fall pretty %ell on a strai!ht line1
# fitted a line to the points 7for the techniEue of linear re!ression0 see note on p1 2238 and then took
the lierty of proHectin! it on into the future1 "ore recently0 Hust as this ook %as !oin! to press0 #
sho%ed this section to Professor )od!kin0 and he told me the most recent data of %hich he %as
a%areA the duckilled platypus !enome0 %hich %as seEuenced in 355; 7the platypus %as a !ood
choice0 ecause of its strate!ic position in the tree of lifeA the ancestor that it shares %ith us lived
2;5 million years a!o0 %hich is nearly three times as lon! a!o as the extinction of the dinosaurs81
#(ve dra%n the platypus(s point as a star on the !raph0 and you can see that it fits pretty %ell near the
proHected line that %as calculated from the earlier data1
The slope of the line for %hat # am no% callin! 7%ithout permission8 )od!kin(s -a% is only
sli!htly shallo%er than that for "oore(s -a%1 The doulin! time is a it more than t%o years0 %here
the "oore(s -a% doulin! time is a it less than t%o years1 '*& technolo!y is intensely dependent
on computers0 so it(s a !ood !uess that )od!kin(s -a% is at least partly dependent on "oore(s -a%1
The arro%s on the ri!ht indicate the !enome siFes of various creatures1 #f you follo% the arro%
to%ards the left until it hits the slopin! line of )od!kin(s -a%0 you can read off an estimate of %hen
it %ill e possile to seEuence a !enome the same siFe as the creature concerned for only PS20555
7of today(s money81 ,or a !enome the siFe of yeast(s0 %e need %ait only till aout 35351 ,or a ne%
mammal !enome 7as far as this kind of ack:of:envelope calculation is concerned0 all mammals are
eEually expensive80 the estimated date is Hust this side of 35>51 #t(s an exhilaratin! prospectA a
massive dataase of '*& seEuences0 cheaply and easily otained from all corners of the animal and
plant kin!doms1 'etailed '*& comparisons %ill fill in all the !aps in our kno%led!e aout the
actual evolutionary relatedness of every species to every otherA %e shall kno%0 %ith complete
certainty0 the entire family tree of all livin! creatures1 L Goodness kno%s ho% %e(ll plot itJ it %on(t
fit on any practical:siFed sheet of paper1
'Ho%'&in's 1a,'
The lar!est:scale attempt in that direction so far has een made y a !roup associated %ith
'avid )illis0 rother of 'anny )illis %ho pioneered one of the first supercomputers1 The )illis plot
makes the tree dia!ram more compact y %rappin! it around in a circle1 6ou can(t see the !ap0
%here the t%o ends almost meet0 ut it lies et%een the (acteria( and the (archaea(1 To see ho% the
circular plot %orks0 look at the !reatly stripped:do%n version tattooed on the ack of 'r Clare
'(&lerto of the $niversity of "elourne0 %hose enthusiasm for Foolo!y is more than skin deep1
Clare has !raciously allo%ed me to reproduce the photo!raph in this ook 7see colour pa!e 3=81 )er
tattoo includes a small sample of ei!hty:six species 7the numer of terminal t%i!s81 6ou can see the
!ap in the circular plot0 and ima!ine the circle opened out1 The smaller numer of illustrations
around the ed!e are strate!ically chosen from acteria0 protoFoa0 plants0 fun!i0 and four animal
phyla1 The verterates are represented y the %eedy sea dra!on on the ri!ht0 a surprisin! fish0
protected y its resemlance to sea%eed1 The )illis circular plot is the same0 except that it has three
thousand species1 Their names appear around the outside ed!e of the circle aove0 far too small to
read : thou!h )omo sapiens is helpfully marked (6ou are here(1 6ou can !et an idea of ho% sparse a
samplin! of the tree even this hu!e plot is %hen # tell you that the closest relatives of humans that it
can fit in the circle are rats and mice1 The mammals had to e stripped do%n drastically0 in order to
fit in all the other ranches of the tree to the same depth1 .ust ima!ine tryin! to plot a similar tree
%ith ten million species instead of the three thousand included here1 &nd ten million is not the most
extrava!ant estimate of the numer of survivin! species1 #t(s %ell %orth do%nloadin! the )illis tree
from his %esite 7see endnotes80 and then printin! it as a %all han!in!0 on a piece of paper %hich0
they recommend0 should e at least => inches %ide 7even i!!er %ould e an advanta!e81
The Hillis #lot
*o%0 %hile %e are talkin! molecules0 %e have some unfinished usiness left over from the
chapter on evolutionary clocks1 There0 %e looked at tree rin!s0 and at various kinds of radioactive
clocks0 ut %e deferred consideration of the so:called molecular clock until %e had learned aout
some other aspects of molecular !enetics1 The time has no% come1 Think of this section as an
appendix to the chapter on clocks1
The molecular clock assumes that evolution is true0 and that it proceeds at a sufficiently
constant rate throu!h !eolo!ical time to e used as a clock in its o%n ri!ht0 provided that it can e
calirated usin! fossils0 %hich are in turn calirated %ith radioactive clocks1 .ust as a candle clock
assumes that candles urn at a fixed and kno%n rate0 and a %ater clock assumes that %ater drains
from a ucket at a rate that can e calirated0 and a !randfather clock assumes that a pendulum
s%in!s at a fixed rate0 so the molecular clock assumes that there are certain aspects of evolution
itself that proceed at a fixed rate1 That fixed rate can e calirated a!ainst those parts of the
evolutionary record that are %ell documented %ith 7radioactively datale8 fossils1 Once calirated0
the molecular clock can then e used for other parts of evolution that are not %ell documented y
fossils1 ,or example0 it can e used for animals that don(t have hard skeletons and seldom fossiliFe1
*ice idea0 ut %hat !ives us the ri!ht to hope that %e can find evolutionary processes that
!o at a fixed rateB #ndeed0 much evidence su!!ests that evolutionary rates are hi!hly variale1 -on!
efore the modern era of molecular iolo!y0 .1 B1 S1 )aldane proposed the darwin as a measure of
evolutionary rates1 Suppose that0 over evolutionary time0 some measured characteristic of an animal
is chan!in! in a consistent direction1 ,or example0 suppose the mean le! len!th is increasin!1 #f0
over a period of a million years0 le! len!th increases y a factor of e 731@2; 1 1 10 a numer chosen
for reasons of mathematical convenience0 %hich %e needn(t !o into80 L the rate of evolutionary
chan!e is said to e one dar%in1 )aldane himself assessed the rate of evolution of the horse as
approximately >5 millidar%ins0 %hile it has een su!!ested that the evolution of domestic animals
under artificial selection should e measured in kilodar%ins1 The rate of evolution of !uppies
transplanted to a predator:free stream0 as descried in Chapter =0 has een estimated as >=
kilodar%ins1 The evolution of (livin! fossils( such as (in$"la 7 pa!e 2>58 is proaly to e measured
in microdar%ins1 6ou !et the pointA rates of evolution of thin!s that you can see and measure0 like
le!s and eaks0 are hu!ely variale1
#f rates of evolution are so variale0 ho% can %e hope to use them as a clockB This is %here
molecular !enetics comes to the rescue1 &t first si!ht0 it %ill not e clear ho% this can e so1 When
measurale characteristics like le! len!th evolve0 %hat %e are seein! is the out%ard and visile
manifestation of an underlyin! !enetic chan!e1 )o%0 then0 can it e the case that rates of chan!e at
the molecular level provide a !ood clock %hile rates of le! or %in! evolution don(tB #f le!s and
eaks under!o chan!e at rates ran!in! from microdar%ins to kilodar%ins0 %hy should molecules e
any more reliale as clocksB The ans%er is that the !enetic chan!es that manifest themselves in
out%ard and visile evolution : of thin!s like le!s and arms : are a very small tip of the iceer!0 and
they are the tip that is heavily influenced y varyin! natural selection1 The maHority of !enetic
chan!e at the molecular level is ne"tral0 and can therefore e expected to proceed at a rate that is
independent of usefulness and mi!ht even e approximately constant %ithin any one !ene1 &
neutral !enetic chan!e has no effect on the survival of the animal0 and this is a helpful credential for
a clock1 This is ecause !enes that affect survival0 positively or ne!atively0 %ould e expected to
evolve at a chan!ed rate0 reflectin! this1
When the neutral theory of molecular evolution %as first proposed y0 amon! others0 the
!reat .apanese !eneticist "otoo Dimura0 it %as controversial1 Some version of it is no% %idely
accepted and0 %ithout !oin! into the detailed evidence here0 # am !oin! to accept it in this ook1
Since # have a reputation as an arch:(adaptationist( 7alle!edly osessed %ith natural selection as the
maHor or even only drivin! force of evolution8 you can have some confidence that if even # support
the neutral theory it is unlikely that many other iolo!ists %ill oppose itKL
& neutral mutation is one that0 althou!h easily measurale y molecular !enetic techniEues0
is not suHect to natural selection0 either positive or ne!ative1 (Pseudo!enes( are neutral for one kind
of reason1 They are !enes that once did somethin! useful ut have no% een sidelined and are never
transcried or translated1 They mi!ht as %ell not exist0 as far as the animal(s %elfare is concerned1
But as far as the scientist is concerned they very much exist0 and they are exactly %hat %e need for
an evolutionary clock1 Pseudo!enes are only one class of those !enes that are never translated in
emryolo!y1 There are other classes %hich are preferred y scientists for molecular clocks0 ut #
%on(t !o into detail1 What pseudo!enes are useful for is emarrassin! creationists1 #t stretches even
their creative in!enuity to make up a convincin! reason %hy an intelli!ent desi!ner should have
created a pseudo!ene : a !ene that does asolutely nothin! and !ives every appearance of ein! a
superannuated version of a !ene that used to do somethin! : unless he %as delierately settin! out
to fool us1
-eavin! pseudo!enes aside0 it is a remarkale fact that the !reater part 79= per cent in the
case of humans8 of the !enome mi!ht as %ell not e there0 for all the difference it makes1 The
neutral theory applies even to many of the !enes in the remainin! = per cent : the !enes that are read
and used1 #t applies even to !enes that are totally vital for survival1 # must e clear here1 We are not
sayin! that a !ene to %hich the neutral theory applies has no effect on the ody1 What %e are sayin!
is that a mutant version of the !ene has exactly the same effect as the unmutated version1 )o%ever
important or unimportant the !ene itself may e0 the mutated version has the same effect as the
unmutated version1 $nlike pseudo!enes0 %here the !ene itself can properly e descried as neutral0
%e are no% talkin! aout cases %here it is only m"tations 7i1e1 chan!es in !enes8 that can strictly e
descried as neutral0 not !enes themselves1
"utations can e neutral for various reasons1 The '*& code is a (de!enerate code(1 This is a
technical term meanin! that some code (%ords( are exact synonyms of each other1L When a !ene
mutates into one of its synonyms0 you mi!ht as %ell not other to call it a mutation at all1 #ndeed0 it
isn(t a mutation0 as far as conseEuences on the ody are concerned1 &nd for the same reason it isn(t a
mutation at all as far as natural selection is concerned1 But it is a mutation as far as molecular
!eneticists are concerned0 for they can see it usin! their methods1 #t is as thou!h # %ere to chan!e
the font in %hich # %rite a %ord0 say kan!aroo to kan!aroo1 6ou can still read the %ord0 and it still
means the same &ustralian hoppin! animal1 The chan!e of typeface from "inion to )elvetica is
detectale ut irrelevant to the meanin!1
*ot all neutral mutations are Euite so neutral as that1 Sometimes the ne% !ene translates into
a different protein0 ut the (active site( 7rememer the carefully shaped (dents( that %e met in Chapter
;8 of the ne% protein remains the same as the old one1 ConseEuently0 there is literally no effect on
the emryonic development of the ody1 The unmutated and the mutated form of the !ene are still
synonyms as far as their effects on odies are concerned1 #t is also possile 7althou!h (ultra:
'ar%inists( like me incline a!ainst the idea8 that some mutations really do chan!e the ody0 ut in
such a %ay as to have no effect on survival0 one %ay or the other1
So0 to sum up on the neutral theory0 to say that a !ene0 or a mutation0 is (neutral( doesn(t
necessarily mean that the !ene itself is useless1 #t could e vitally important to the animal(s survival1
What it means is that the mutated form of a !ene : %hich mi!ht or mi!ht not e important for
survival : is no different from the unmutated form %ith respect to its effects 7%hich mi!ht e very
important8 on survival1 &s it happens0 it is proaly true to say that most mutations are neutral1
They are undetectale y natural selection0 ut detectale y molecular !eneticistsJ and that is an
ideal comination for an evolutionary clock1
*one of this is to do%n!rade the all:important tip of the iceer! : the minority of mutations
that are not neutral1 #t is they that are selected0 positively or ne!atively0 in the evolution of
improvements1 They are the ones %hose effects %e actually see : and natural selection (sees( too1
They are the ones %hose selection !ives livin! thin!s their reathtakin! illusion of desi!n1 But it is
the rest of the iceer! : the neutral mutations0 %hich are in the maHority : that concern us %hen %e
are talkin! aout the molecular clock1
&s !eolo!ical time !oes y0 the !enome is suHected to a rain of attrition in the form of
mutations1 #n that small portion of the !enome %here the mutations really matter for survival0
natural selection soon !ets rid of the ad ones and favours the !ood ones1 The neutral mutations0 on
the other hand0 simply pile up0 unpunished and unnoticed : except y molecular !eneticists1 &nd
no% %e need a ne% technical termA fixation1 & ne% mutation0 if it is !enuinely ne%0 %ill have a lo%
freEuency in the !ene pool1 #f you revisit the !ene pool a million years later0 it is possile that the
mutation %ill have increased in freEuency to 255 per cent or somethin! close to it1 #f that happens0
the mutation is said to have (!one to fixation(1 We shall no lon!er think of it as a mutation1 #t has
ecome the norm1 The ovious %ay for a mutation to !o to fixation is for natural selection to favour
it1 But there is another %ay1 #t can !o to fixation y chance1 .ust as a once proud surname can die
out for lack of male heirs0 so the alternatives to the mutation %e are talkin! aout can Hust happen to
disappear from the !ene pool1 The mutation itself can ecome freEuent in the !ene pool0 y the
same luck as has led (Smith( to emer!e as the commonest surname in En!land1 Of course it is much
more interestin! if the !ene !oes to fixation for a !ood reason : that(s natural selection : ut it can
also happen y chance0 !iven a lar!e enou!h numer of !enerations1 &nd !eolo!ical time is vast
enou!h for neutral mutations to !o to fixation at a predictale rate1 The rate at %hich they do so
varies0 ut it is characteristic of particular !enes0 and0 !iven that most mutations are neutral0 this is
precisely %hat makes the molecular clock possile1
#t(s fixation that matters for the molecular clock0 ecause (fixed( !enes are the ones that %e
look at %hen %e compare t%o modern animals to try to estimate ho% lon! a!o their ancestors split
apart1 ,ixed !enes are the !enes that characteriFe a species1 They are the ones that are all ut
universal in the !ene pool1 &nd %e can compare the !enes that have ecome fixed in one species
%ith the !enes that have ecome fixed in another0 in order to estimate ho% recently the t%o species
split apart1 There are complications0 %hich # %on(t !o into ecause 6an Won! and # discussed them
fully in (The Epilo!ue to the +elvet Worm(s Tale(1 With reservations0 and %ith various important
correction factors0 the molecular clock %orks1
.ust as radioactive clocks tick at hu!ely variale speeds0 %ith half:lives ran!in! from
fractions of a second throu!h to tens of illions of years0 so different !enes provide a marvellous
spread of molecular clocks0 suitale for timin! evolutionary chan!e on scales ran!in! from a
million to a illion years0 and all sta!es in et%een1 .ust as each radioactive isotope has its
characteristic half:life0 so each !ene has a characteristic turnover rate : the rate at %hich ne%
mutations typically !o to fixation y random chance1 )istone !enes characteristically turn over at a
rate of one mutation per illion years1 ,irinopeptide !enes are a thousand times faster0 %ith a
turnover of one ne% mutation fixed per million years1 Cytochrome:C and the suite of haemo!loin
!enes have intermediate turnovers0 %ith times to fixation measured in millions to tens of millions of
*either radioactive clocks nor molecular clocks tick in a re!ular fashion like a pendulum
clock or a %atch1 #f you could hear them tickin!0 they(d sound like a Gei!er counter0 the radioactive
clocks literally so since a Gei!er counter is precisely %hat you %ould use to listen to them1 &
Gei!er counter doesn(t tick re!ularly0 like a %atchJ it ticks at random0 the ticks comin! in stran!e0
stutterin! ursts1 That(s ho% mutations0 and fixations0 %ould sound0 if %e could hear them on the
immensely lon! timescale of !eolo!y1 But0 %hether stutterin! like a Gei!er counter or tickin!
metronomically like a %atch0 the important thin! aout a timekeeper is that it should tick at a
kno%n aera$e rate1 That(s %hat radioactive clocks do0 and that(s %hat molecular clocks do1
# introduced the molecular clock y sayin! that it assumes the fact of evolution and therefore
can(t e used in evidence of it1 But no%0 havin! understood ho% the clock %orks0 %e can see that #
%as too pessimistic1 The very existence of pseudo!enes : useless0 untranscried !enes that ear a
marked resemlance to useful !enes : is a perfect example of the %ay animals and plants have their
history %ritten all over them1 But that is a topic that must %ait for the next chapter1
L 6ou may e surprised to hear that horses evolved in *orth &merica0 ecause it is %idely
kno%n that %hen the European invaders first came to the &mericas0 the si!ht of them on horseack
amaFed the natives1 The ulk of horse evolution did indeed take place in &merica1 Then horses
spread to the rest of the %orld0 shortly 7y !eolo!ical standards8 efore !oin! extinct in &merica1
They are &merican animals that have een re:introduced to &merica y man1
L & sin!le one in mammals1 The reptilian lo%er Ha% is more complicated : and therey
han!s a fascinatin! tale that # reluctantly omitted from this ook 7you can(t have everythin!81 #n an
amaFin! feat of evolutionary le!erdemain0 the smaller ones of the reptilian lo%er Ha% %ere
coopted into the mammalian ear0 %here they constitute an exEuisitely delicate rid!e0 to transport
sound from the eardrum to the inner ear1
L The 'utch (%ildeeest( is increasin!ly used in preference to (!nu(1 # am tryin! to save (!nu(
ecause0 if it dies out alto!ether0 the %itty son! y ,landers and S%ann %on(t make sense any more1
7(Gnor am # in the least G -ike that dreadful harteeest G Oh !no !no !no0 #(m a !nuK(8
L # presume my readers kno% etter than the author7s8 of -eviticus0 %ho thou!ht that ats
%ere irds1 #n chapter 220 verses 24:290 is a lon! list of irds that are an aomination0 e!innin!
%ith the ea!le and endin! %ith (the stork0 the heron after her kind0 and the lap%in!0 and the at(1 #t is
a separate Euestion %hy it %as necessary to condemn any animals as aominations1 #t %as a
common practice in many reli!ions1
L Biolo!ists used to cite plant haemo!loin as a possile example of '*& orro%in! y
plants from the animal kin!dom1 Plants of the pea family 7-e!uminosae8 have on their roots
(nodules(0 in %hich d%ell acteria that capture nitro!en from the atmosphere and make it availale
to the plants1 This is %hy farmers often include a le!uminous crop0 such as clover or a vetch0 in
their rotation1 #t puts valuale nitro!en in the soil0 especially if the clover crop is plou!hed under1
The nodules are a reddish colour ecause they contain a form of haemo!loin0 similar to the
oxy!en:transportin! molecule that makes our lood red1 The !enes for makin! haemo!loin are in
the plant !enome0 not the acterial !enome1 )aemo!loin is important to the acteria0 %hich need
oxy!en0 and it can e re!arded as part of the deal et%een acteria and plantsA the acteria !ive the
plants usale nitro!en0 %hile the plants !ive the acteria a house0 and usale oxy!en delivered via
haemo!loin1 Since %e are accustomed to associatin! haemo!loin %ith lood0 it %as natural to
%onder %hether a !ene for makin! it had someho% een (orro%ed( from an animal !enome0
perhaps ferried y a acterium1 #t %ould0 indeed0 have een a very valuale idea to (orro%(1
$nfortunately for this appealin! idea : the ultimate lood transfusion : molecular iolo!ical
evidence sho%s that haemo!loins are ancient deniFens of plant !enomes1 They are not orro%ed1
They have een there from ancient times1
L #t is a little:kno%n fact that some dinosaurs had a !an!lion in the pelvis0 %hich %as so
lar!e 7at least relative to the rain in the head8 as almost to deserve the title of second rain1 This
prompted the follo%in! deli!htfully %itty verse y the &merican comic %riter Bert -eston Taylor
Behold the mi!hty dinosaur0
,amous in prehistoric lore0
*ot only for his po%er and stren!th
But for his intellectual len!th1
6ou %ill oserve y these remains
The creature had t%o sets of rains :
One in his head 7the usual place80
The other at his spinal ase0
Thus he could reason A priori
&s %ell as A posteriori1
*o prolem othered him a it
)e made oth head and tail of it1
So %ise %as he0 so %ise and solemn0
Each thou!ht filled Hust a spinal column1
#f one rain found the pressure stron!
#t passed a fe% ideas alon!1
#f somethin! slipped his for%ard mind
(T%as rescued y the one ehind1
&nd if in error he %as cau!ht
)e had a savin! afterthou!ht1
&s he thou!ht t%ice efore he spoke
)e had no Hud!ment to revoke1
Thus he could think %ithout con!estion
$pon oth sides of every Euestion1
Oh0 !aFe upon this model east0
'efunct ten million years at least1
L '(&rcy Thompson %as surely one of the most erudite scientists ever1 *ot only did he %rite
famously eautiful En!lish of a patrician character0 not only %as he a pulished mathematician and
classical scholar as %ell as Professor of *atural )istory at Scotland(s oldest university0 ut his ook
is laced %ith Euotations0 %hich he assumed he had no need to translate 7ho% times have chan!ed80
in -atin0 Greek0 #talian0 German0 ,rench and even Provencal 7the last he did dei!n to translate : into
L Strictly speakin!0 t%o shapes are homeomorphic if you can distort one to ecome the other
%ithout reakin! it0 and %ithout any ne% touchin!s1
L Perhaps (all livin! creatures( calls for a note of caution1 #n an earlier section of this chapter
%e sa% that %hile the principle of (*o orro%in!( is almost completely appropriate for animals and
plants0 acteria are different1 &mon! acteria 7and archaea0 %hich are superficially like acteria ut
rather distantly related8 there is plenty of sharin! of !enes1 Whereas animals use sexual couplin! to
exchan!e '*& %ithin species0 acteria use their o%n form of (Copy and Paste( to pass '*&
around0 even et%een distantly related species1 &lthou!h # %as ri!ht to extol the (one true tree of
life( for animals and plants0 the %hole usiness !ets messier %hen %e turn to microor!anisms1 &s
my collea!ue the philosopher 'an 'ennett has put it0 %here the tree of life for animals is a
maHestically spreadin! oak0 that for acteria is more like a anyan1 Where acteria are concerned0
there is somethin! to e said for compilin! a (one true tree( for each !ene separately0 re!ardless of
%hich particular kinds of acteria it happens to e travellin! around in1 What a fascinatin! prospect1
)o% 'ar%in %ould have loved it1
L When # first read #alc"l"s %ade Easy y Silvanus P1 Thompson0 on the recommendation
of my en!ineer !randfather0 it !ave me !ooseumps %hen Thompson introduced e in italics as ( a
n"mber neer to be for$otten(1 One conseEuence of usin! e rather than0 say0 30 as the factor of
choice0 is that you can calculate the dar%ins directly y sutractin! natural lo!arithms from one
another1 Other scientists have proposed0 as a unit of evolutionary rate0 the haldane1
L # have even een called an (ultra:'ar%inist(0 a !ie that # find less insultin! than its coiners
perhaps intended1
L ('e!enerate( is not the same 7thou!h the t%o terms are often confused8 as (redundant(0
another technical term of #nformation Theory1 & redundant code is one in %hich the same messa!e
is conveyed more than once 7e1!1 (She is a female %oman( conveys the messa!e of her sex three
times81 Redundancy is used y en!ineers to !uard a!ainst transmission errors1 & de!enerate code is
one in %hich more than one (%ord( is used to mean the same thin!1 #n the !enetic code0 for example0
C$C and C$G oth spell (-eucine(A a mutation from C$C to C$G therefore makes no difference1
That(s (de!enerate(1
C)&PTER 22
)#STOR6 WR#TTE* &-- O+ER $S
# BEG&* this ook y ima!inin! a teacher of -atin forced to %aste time and ener!y
defendin! the proposition that the Romans and their lan!ua!e ever existed1 -et(s return to that
thou!ht0 and ask %hat actually is the evidence for the Roman Empire and the -atin lan!ua!e1 # live
in Britain %here0 as in the rest of Europe0 Rome %rote her si!nature all over the map0 carved her
%ays into the landscape0 %ove her lan!ua!e throu!h ours and her history throu!h our literature1
Walk the len!th of )adrian(s Wall0 %hose preferred local name is still (The Roman Wall(1 Walk0 as #
%alked Sunday after Sunday in crocodile formation from my oardin! school in 7relatively8 ne%
Salisury0 to the Roman flint fort of Old Sarum0 and commune %ith the ima!ined !hosts of dead
le!ions1 $nfold an Ordnance Survey map of En!land1 Wherever you see a lon!0 dead strai!ht
country road0 especially %hen there are !reen field !aps et%een stretches of road or cart track that
you can exactly line up %ith a ruler0 you(ll almost al%ays find a Roman lael eside it1 +esti!es of
the Roman Empire are all around us1
-ivin! odies0 too0 have their history %ritten all over them1 They ristle %ith the iolo!ical
eEuivalent of Roman roads0 %alls0 monuments0 potsherds0 even ancient inscriptions carved into the
livin! '*&0 ready to e deciphered y scholars1
BristleB 6es0 literally1 When you are cold0 or adly fri!htened0 or haunted y the peerless
craftsmanship of a Shakespeare sonnet0 you !et !ooseumps1 WhyB Because your ancestors %ere
normal mammals %ith hairs all over0 and these %ere raised or lo%ered at the ehest of sensitive
odily thermostats1 Too cold0 and the hairs %ere erected to plump up the layer of insulatin! trapped
air1 Too %arm0 and the coat %as flattened to allo% ody heat to escape more easily1 #n later
evolution0 the hair:erection system %as hiHacked for social communication purposes0 and ecame
involved in The Expression of the Emotions0 as 'ar%in %as one of the first to appreciate in his ook
of that name1 # can(t resist sharin! %ith you some lines : vinta!e 'ar%in : from that ookA
"r Sutton0 the intelli!ent keeper in the Ioolo!ical Gardens0 carefully oserved for me the
ChimpanFee and Oran!J and he states that %hen they are suddenly fri!htened0 as y a thunderstorm0
or %hen they are made an!ry0 as y ein! teased0 their hair ecomes erect1 # sa% a chimpanFee %ho
%as alarmed at the si!ht of a lack coalheaver0 and the hair rose all over his ody 1 1 1 # took a
stuffed snake into the monkey:house0 and the hair on several of the species instantly ecame erect 1 1
1 When # sho%ed a stuffed snake to a Peccary0 the hair rose in a %onderful manner alon! its ackJ
and so it does %ith a %ild oar %hen enra!ed1
The hackles are raised in an!er1 #n fear also0 hairs stand on end to increase the ody(s
apparent siFe and scare off dan!erous rivals or predators1 Even %e naked apes still have the
machinery to raise non:existent 7or arely:existent8 hairs0 and %e call it !ooseumps1 The hair:
erection machinery is a esti$e0 a non:functional relic of somethin! that did a useful Ho in our lon!:
dead ancestors1 +esti!ial hairs are amon! the many instances of history %ritten all over us1 They
constitute persuasive evidence that evolution has occurred0 and a!ain it comes not from fossils ut
from modern animals1
&s %e sa% in the previous chapter0 %here # compared it to a comparaly siFed fish such as a
dorado0 you don(t have to di! very deep inside a dolphin to uncover its history of life on dry land1
'espite its streamlined0 fish:like exterior0 and despite the fact that it no% makes its entire livin! in
the sea and %ould soon die if eached0 a dolphin0 ut not a dorado0 has (land mammal( %oven
throu!h its very %arp and %oof1 #t has lun!s not !ills0 and %ill dro%n like any land animal if
prevented from comin! up for air0 althou!h it can hold its reath for much lon!er than a land
mammal1 #n all sorts of %ays0 its air:reathin! apparatus is chan!ed to fit its %atery %orld1 #nstead
of reathin! throu!h t%o little nostrils at the end of its nose like any normal land mammal0 it has a
sin!le nostril in the top of its head0 %hich enales it to reathe %hile only Hust reakin! the surface1
This (lo%hole( has a ti!ht:sealin! valve to keep %ater out0 and a %ide ore to minimiFe the time
needed for reathin!1 #n an 2;>= communication to the Royal Society0 %hich 'ar%in0 as a ,ello%0
%ould Euite likely have read0 ,rancis Sison EsE1 L %roteA (The muscles that open and close the
lo%:hole0 and that act upon the various sacs0 form one of the most complicated yet most
exEuisitely adHusted pieces of machinery that either nature or art presents1( The dolphin(s lo%hole
!oes to !reat len!ths to correct a prolem that %ould never have arisen at all if only it reathed %ith
!ills0 like a fish1 &nd many of the details of the lo%hole can e seen as corrections to secondary
prolems that arose %hen the air intake mi!rated from the nostrils to the top of the head1 & real
desi!ner %ould have planned it in the top of the head in the first place : that(s if he hadn(t decided to
aolish lun!s and !o for !ills any%ay1 Throu!hout this chapter0 %e shall continually find examples
of evolution correctin! an initial (mistake( or historical relic y post hoc compensation or t%eakin!0
rather than y !oin! ack to the dra%in! oard as a real desi!ner %ould1 #n any case0 the elaorate
and complex !ate%ay to the lo%hole is eloEuent testimony to the dolphin(s remote ancestry on dry
#n countless other %ays0 dolphins and %hales could e said to have their ancient history
%ritten all over and throu!h them0 like vesti!es of Roman roads dra%n out in dead strai!ht cart
tracks and ridle%ays across the map of En!land1 Whales have no hind le!s0 ut there are tiny
ones0 uried deep inside them0 %hich are the remnants of the pelvic !irdle and hind le!s of their
lon!:!one %alkin! ancestors1 The same is true of the sirenians or sea co%s 7#(ve already mentioned
them several timesA the manatees0 du!on!s and the ;:yard:lon! Steller(s sea co%0 hunted to
extinction y humans81 L Sirenians are very different from %hales and dolphins0 ut they are the
only other !roup of %holly marine mammals that never come ashore1 Where dolphins are fast0
actively intelli!ent carnivores0 manatees and du!on!s are slo%0 dreamy herivores1 &t the manatee
aEuarium that # visited in %estern ,lorida0 for once # didn(t ra!e a!ainst the loudspeakers playin!
music1 #t %as sleepy la!oon music and it seemed so lan!uidly appropriate that all %as for!iven1
"anatees and du!on!s float effortlessly in hydrostatic eEuilirium0 not y means of a s%im ladder
as fish do 7see elo%80 ut throu!h ein! eEuipped %ith heavy ones as a counter%ei!ht to the
natural uoyancy of their luer1 Their specific !ravity is therefore very close to that of %ater0 and
they can make fine adHustments to it y pullin! in or expandin! the ri ca!e1 The precision of their
uoyancy control is enhanced y the possession of a separate cavity for each lun!A they have t%o
independent diaphra!ms1
'olphins and %hales0 du!on!s and manatees !ive irth to live aies0 like all mammals1
That hait is not actually peculiar to mammals1 "any fish are liveearers0 ut they do it in a very
different %ay 7actually a fascinatin! variety of very different %ays0 doutless independently
evolved81 The dolphin(s placenta is unmistakaly mammalian0 and so is its hait of sucklin! the
youn! %ith milk1 #ts rain is also eyond Euestion the rain of a mammal0 and a very advanced
mammal at that1 The cereral cortex of a mammal is a sheet of !rey matter0 %rapped around the
outside of the rain1 Gettin! rainier partly consists in increasin! the area of the sheet1 This could
e done y increasin! the total siFe of the rain0 and of the skull that houses it1 But there are
do%nsides to havin! a i! skull1 #t makes it harder to e orn0 for one thin!1 &s a result0 rainy
mammals contrive to increase the area of the sheet %hile stayin! %ithin limits set y the skull0 and
they do it y thro%in! the %hole sheet into deep folds and fissures1 This is %hy the human rain
looks like a %rinkled %alnutJ and the rains of dolphins and %hales are the only ones to rival those
of us apes for %rinkliness1 ,ish rains don(t have %rinkles at all1 #ndeed0 they don(t have a cereral
cortex0 and the %hole rain is tiny compared to a dolphin(s or human(s1 The dolphin(s mammalian
history is deeply etched into the %rinkled surface of its rain1 #t(s a part of its mammalness0 alon!
%ith the placenta0 milk0 a four:chamered heart0 a lo%er Ha% havin! only a sin!le one0 %arm:
loodedness0 and many other specifically mammalian features1
!rains o$ h("an ?to#@, %ol#hin ?"i%%le@, *ro,n tro(t ?*otto"@ ?not to sale@
Warm:looded is %hat %e call mammals and irds0 ut really %hat they have is the aility
to keep their temperature constant0 re!ardless of the outside temperature1 This is a !ood idea0
ecause the chemical reactions in a cell can then all e optimiFed to %ork est at a particular
temperature1 (Cold:looded( animals are not necessarily cold1 & liFard has %armer lood than a
mammal if oth happen to e out in the midday sun in the Sahara desert1 & liFard has colder lood
than a mammal if they are out in the sno%1 The mammal has the same temperature all the time0 and
it has to %ork hard to keep it constant0 usin! internal mechanisms1 -iFards use external means to
re!ulate their temperature0 movin! into the sun %hen they need to %arm themselves up0 and into the
shade %hen they need to cool do%n1 "ammals re!ulate their ody temperature more accurately0
and dolphins are no exception1 Once a!ain0 their mammal history is %ritten all over them0 even
thou!h they have reverted to life in the sea0 %here most creatures don(t maintain a constant
The odies of %hales and sirenians aound in historical relics that %e notice ecause they
live in a very different environment from their land:d%ellin! ancestors1 & similar principle applies
to irds that have lost the hait and eEuipment of fli!ht1 *ot all irds fly0 ut all irds carry at least
relics of the apparatus of fli!ht1 Ostriches and emus are fast runners that never fly0 ut they have
stus of %in!s as a le!acy from remote flyin! ancestors1 Ostrich %in! stus0 moreover0 have not
completely lost their usefulness1 &lthou!h much too small to fly %ith0 they seem to have some sort
of alancin! and steerin! role in runnin!0 and they enter into social and sexual displays1 Di%i %in!s
are too small to e seen outside the ird(s fine coat of feathers0 ut vesti!es of %in! ones are there1
"oas lost their %in!s entirely1 Their home country of *e% Iealand0 y the %ay0 has more than its
fair share of fli!htless irds0 proaly ecause the asence of mammals left %ide open niches to e
filled y any creature that could !et there y flyin!1 But those flyin! pioneers0 havin! arrived on
%in!s0 later lost them as they filled the vacant mammal roles on the !round1 This proaly doesn(t
apply to the moas themselves0 %hose ancestors0 as it happened0 %ere already fli!htless efore the
!reat southern continent of Gond%ana roke up into fra!ments0 *e% Iealand amon! them0 each
earin! its o%n car!o of Gond%anan animals1 #t surely does apply to kakapos0 *e% Iealand(s
fli!htless parrots0 %hose flyin! ancestors apparently lived so recently that kakapos still try to fly
althou!h they lack the eEuipment to succeed1 #n the %ords of the immortal 'ou!las &dams0 in (ast
#hance to See0
#t is an extremely fat ird1 & !ood:siFed adult %ill %ei!h aout six or seven pounds0 and its
%in!s are Hust aout !ood for %i!!lin! aout a it if it thinks it(s aout to trip over somethin! : ut
flyin! is completely out of the Euestion1 Sadly0 ho%ever0 it seems that not only has the kakapo
for!otten ho% to fly0 ut it has also for!otten that it has for!otten ho% to fly1 &pparently a seriously
%orried kakapo %ill sometimes run up a tree and Hump out of it0 %hereupon it flies like a rick and
lands in a !raceless heap on the !round1
While ostriches0 emus and rheas are !reat runners0 pen!uins and Galapa!os fli!htless
cormorants are !reat s%immers1 # %as privile!ed to s%im %ith a fli!htless cormorant in a lar!e rock
pool on the island of #saela0 and # %as enchanted to %itness the speed and a!ility %ith %hich it
sou!ht out one undersea crevice after another0 stayin! under for a reathtakin!ly lon! time 7# had
the advanta!e of a snorkel81 $nlike pen!uins0 %ho use their short %in!s to (fly under%ater(0
Galapa!os cormorants propel themselves %ith their po%erful le!s and hu!e %eed feet0 usin! their
%in!s only as stailiFers1 But all fli!htless irds0 includin! ostriches and their kind0 %hich lost their
%in!s a very lon! time a!o0 are clearly descended from ancestors that used them to fly1 *o
reasonale oserver could seriously dout the truth of that0 %hich means that anyody %ho thinks
aout it should find it very hard : %hy not impossileB : to dout the fact of evolution1
*umerous different !roups of insects0 too0 have lost their %in!s0 or !reatly reduced them1
$nlike primitively %in!less insects such as silverfish0 fleas and lice have lost the %in!s their
ancestors once had1 ,emale !ypsy moths have underdeveloped %in! muscles and don(t fly1 They
don(t need to0 for the males fly to them0 attracted y a chemical lure %hich they can detect at
astoundin! dilutions1 #f the females %ere to move as %ell as the males0 the system proaly
%ouldn(t %ork0 for y the time the male had flo%n up the slo%ly driftin! chemical !radient0 its
source %ould have moved onK
$nlike most insects0 %hich have four %in!s0 the flies0 as their -atin name 'iptera su!!ests0
have only t%o1 The second pair of %in!s has ecome reduced to a pair of (halteres(1 These s%in!
aout like very hi!h:speed #ndian clus0 %hich they resemle0 functionin! as tiny !yroscopes1 )o%
do %e kno% that halteres are descended from ancestral %in!sB Several reasons1 They occupy
exactly the same place in the third se!ment of the thorax as the flyin! %in! occupies in the second
thoracic se!ment 7and the third too in other insects81 They move in the same fi!ure:of:ei!ht pattern
as the %in!s of flies1 They have the same emryolo!y as %in!s and0 althou!h they are tiny0 if you
look at them carefully0 especially durin! development0 you can see that they are stunted %in!s0
clearly modified : unless you are an evolution:denier : from ancestral %in!s1 Testifyin! to the same
story0 there are mutant fruit flies0 so:called homeotic mutants0 %hose emryolo!y is anormal and
%ho !ro% not halteres ut a second pair of %in!s0 like a ee or any other kind of insect1
Halteres on a rane$ly
What %ould the intermediate sta!es et%een %in!s and halteres have looked like0 and %hy
%ould natural selection have favoured the intermediatesB What is the use of half a haltereB .1 W1 S1
Prin!le0 my old Oxford professor %hose foriddin! mien and stiff earin! earned him the nickname
(-au!hin! .ohn(0 %as mainly responsile for %orkin! out ho% halteres %ork1 )e pointed out that all
insect %in!s have tiny sense or!ans in the ase0 %hich detect t%istin! and other forces1 The sense
or!ans at the ase of halteres are very similar : another piece of evidence that halteres are modified
%in!s1 -on! efore halteres evolved0 the information streamin! into the nervous system from the
sense or!ans at their ase %ould enale fast uFFin! %in!s0 %hile flyin!0 to act as rudimentary
!yroscopes1 To the extent that any flyin! machine is naturally unstale0 it needs to compensate %ith
sophisticated instrumentation0 for example !yroscopes1
The %hole Euestion of the evolution of stale and unstale fliers is very interestin!1 -ook at
these t%o pterosaurs0 extinct flyin! reptiles0 contemporaries of the dinosaurs1 &ny aero:en!ineer
could tell you that Rhamphorhynch"s0 the early pterosaur at the top of the picture0 must have een a
stale flier0 ecause of its lon! tail %ith the pin!:pon! at on the end1 Rhamphorhynch"s %ould not
have needed sophisticated !yro:control0 such as flies have %ith their halteres0 ecause its tail made
it inherently stale1 On the other hand0 as the same en!ineer could tell you0 it %ould not have een
very manoeuvrale1 #n any flyin! machine0 there is a trade:off et%een staility and
manoeuvraility1 The !reat .ohn "aynard Smith0 %ho %orked as an aircraft desi!ner efore
returnin! to university to read Foolo!y 7on the !rounds that aeroplanes %ere noisy and old:
fashioned80 pointed out that flyin! animals can move in evolutionary time0 ack and forth alon! the
spectrum of this trade:off0 sometimes losin! inherent staility in the interests of increased
manoeuvraility0 ut payin! for it in the form of increased instrumentation and computation
capaility : rain po%er1 &t the ottom of the picture on the previous pa!e is Anhan$"era0 a late
pterodactyl from the Cretaceous era0 some <5 million years after the .urassic Rhamphorhynch"s1
Anhan$"era had almost no tail at all0 like a modern at1 -ike a at0 it %ould surely have een an
unstale aircraft0 reliant on instrumentation and computation to exercise sutle0 moment:to:moment
control over its fli!ht surfaces1
%hamphorhynchus ?to#@ an% Anhanguera ?*otto"@
Anhan$"era didn(t have halteres0 of course1 #t %ould have used other sense or!ans to
provide the eEuivalent information0 proaly the semicircular canals of the inner ear1 These %ere
indeed very lar!e in those pterosaurs that have een looked at : althou!h0 a touch disappointin!ly
for the "aynard Smith hypothesis0 they %ere lar!e in Rhamphorhynch"s as %ell as Anhan$"era1
But0 to return to the flies0 Prin!le su!!ests that the four:%in!ed ancestors of flies proaly had lon!
adomens0 %hich %ould have made them stale1 &ll four %in!s %ould have acted as rudimentary
!yroscopes1 Then0 he su!!ests0 the ancestors of flies started to move alon! the staility continuum0
ecomin! more manoeuvrale and less stale as the adomen !ot shorter1 The hind %in!s started to
shift more to%ards the !yroscopic function 7%hich they had al%ays performed0 in a small %ay0 as
%in!s80 ecomin! smaller0 and heavier for their siFe0 %hile the fore%in!s enlar!ed to take over
more of the flyin!1 There %ould have een a !radual continuum of chan!e0 as the fore%in!s
assumed ever more of the urden of aviation0 %hile the hind %in!s shrank to take over the avionics1
Worker ants have lost their %in!s0 ut not the capacity to !ro% %in!s1 Their %in!ed history
still lurks %ithin them1 We kno% this ecause Eueen ants 7and males8 have %in!s0 and %orkers are
females %ho could have een Eueens ut %ho0 for environmental0 not !enetic0 reasons failed to
ecome Eueens1L Presumaly %orker ants lost their %in!s in evolution ecause they are a nuisance
and !et in the %ay under!round1 Poi!nant testimony to this is provided y Eueen ants0 %ho use their
%in!s once only0 to fly out of the natal nest0 find a mate0 and then settle do%n to di! a hole for a
ne% nest1 &s they e!in their ne% life under!round the first thin! they do is lose their %in!s0 in
some cases y literally itin! them offA painful 7perhapsJ %ho kno%sB8 evidence that %in!s are a
nuisance under!round1 *o %onder %orker ants never !ro% %in!s in the first place1
7arasiti $ly $ro" the 7hori%ae $a"ily
Proaly for similar reasons0 ants( nests0 and termites( nests0 are home to a horde of %in!less
han!ers:on of many different types0 feedin! on the rich pickin!s s%ept in y the ever:rustlin!
streams of returnin! fora!ers1 &nd %in!s are Hust as much of a hindrance to them as they are to the
ants themselves1 Who %ould ever elieve that the monstrosity on the ri!ht is a flyB 6et %e kno%
from a careful and detailed study of its anatomy that not only is it a fly0 this parasite of termite nests
elon!s to a particular family of flies0 the Phoridae1 On the next pa!e is a more normal memer of
the same family0 %hich presumaly some%hat resemles the %in!ed ancestors of the %eirdly
%in!less creature aove0 althou!h it too is a parasite of social insects : ees in this case1 6ou can
see the similarity to the sickle:shaped head of the %eird monster on the previous pa!e1 &nd the
monster(s stunted %in!s are Hust visile as the tiny trian!les on either side1
Another $ly $ro" the 7hori%ae $a"ily
There is an additional reason for %in!lessness in this riff:raff of lurkers and sEuatters in ants(
and termites( nests1 "any of them 7not the Phorid flies8 have over evolutionary time assumed a
protective resemlance to ants0 either 7or oth8 to fool the ants or to fool %ould:e predators %ho
mi!ht other%ise pick them out from amon! the less palatale and etter:protected ants1 Who0 on
takin! only a casual !lance0 %ould notice that the insect elo%0 %hich lives in ants( nests0 is not an
ant at all ut a eetleB Once a!ain0 ho% do %e kno%B ,rom deep and detailed resemlances to
eetles0 %hich hu!ely outnumer the superficial features in %hich the insect resemles an antA
exactly the same %ay as %e kno% that a dolphin is a mammal and not a fish1 This creature has its
eetle ancestry %ritten all throu!h it0 except 7a!ain as %ith dolphins8 in those features that define its
superficial appearance0 such as its %in!lessness and its ant:like profile1
!eetle %is'(ise% as an ant
.ust as ants and their suterranean fello%:travellers lose their %in!s under!round0 so
numerous different kinds of animals that live in the depths of dark caves %here there is no li!ht
have reduced or lost their eyes0 and are0 as 'ar%in himself noted0 more or less completely lind1
The %ord (tro!loite(L has een coined for an animal that lives only in the darkest part of caves and
is so specialiFed that it can live no%here else1 Tro!loites include salamanders0 fish0 shrimps0
crayfish0 millipedes0 spiders0 crickets and many other animals1 They are very often %hite0 havin!
lost all pi!ment0 and lind1 They usually0 ho%ever0 retain vesti!es of eyes0 and that is the point of
mentionin! them here1 +esti!ial eyes are evidence of evolution1 Given that a cave salamander lives
in perpetual darkness so has no use for eyes0 %hy %ould a divine creator nevertheless furnish it %ith
dummy eyes0 clearly related to eyes ut non:functionalB
Evolutionists0 on their side0 need to come up %ith an explanation for the loss of eyes %here
they are no lon!er needed1 Why not0 it mi!ht e said0 simply han! on to your eyes0 even if you
never use themB "i!ht they not come in handy at some point in the futureB Why (other( to !et rid
of themB *otice0 y the %ay0 ho% hard it is to resist the lan!ua!e of intention0 purpose and
personification1 Strictly speakin!0 # should not have used the %ord (other(0 should #B # should have
said somethin! like0 ()o% does losin! its eyes enefit an individual cave salamander so that it is
more likely to survive and reproduce than a rival salamander that keeps a perfect pair of eyes0 even
thou!h it never uses themB(
Well0 eyes are almost certainly not cost:free1 Settin! aside the ar!ualy modest economic
costs of makin! an eye0 a moist eye socket0 %hich has to e open to the %orld to accommodate the
s%ivellin! eyeall %ith its transparent surface0 mi!ht e vulnerale to infection1 So a cave
salamander that sealed up its eyes ehind tou!h ody skin mi!ht survive etter than a rival
individual that kept its eyes1
But there is another %ay to ans%er the Euestion and0 instructively0 it doesn(t invoke the
lan!ua!e of advanta!e at all0 let alone purpose or personification1 When %e are talkin! aout
natural selection0 %e think in terms of rare eneficial mutations turnin! up and ein! positively
favoured y selection1 But most mutations are disadvanta!eous0 if only ecause they are random
and there are many more %ays of !ettin! %orse than there are %ays of !ettin! etter1L *atural
selection promptly penaliFes the ad mutations1 #ndividuals possessin! them are more likely to die
and less likely to reproduce0 and this automatically removes the mutations from the !ene pool1
Every animal and plant !enome is suHect to a constant omardment of deleterious mutationsA a
hailstorm of attrition1 #t is a it like the moon(s surface0 %hich ecomes increasin!ly pitted %ith
craters due to the steady omardment of meteorites1 With rare exceptions0 every time a !ene
concerned %ith an eye0 for example0 is hit y a maraudin! mutation0 the eye ecomes a little less
functional0 a little less capale of seein!0 a little less %orthy of the name of eye1 #n an animal that
lives in the li!ht and uses the sense of si!ht0 such deleterious mutations 7the maHority8 are s%iftly
removed from the !ene pool y natural selection1
But in total darkness the deleterious mutations that omard the !enes for makin! eyes are
not penaliFed1 +ision is impossile any%ay1 The eye of a cave salamander is like the moon0 pitted
%ith mutational craters that are never removed1 The eye of a dayli!ht:d%ellin! salamander is like
the Earth0 hit y mutations at the same rate as cave:d%ellers( eyes0 ut %ith each deleterious
mutation 7crater8 ein! cleaned off y natural selection 7erosion81 Of course0 the story of the cave:
d%eller(s eye isn(t only a ne!ative oneA positive selection comes in too0 to favour the !ro%th of
protective skin over the vulnerale sockets of the optically deterioratin! eyes1
&mon! the most interestin! of historical relics are those features that are used for somethin!
7and so are not vesti!es in the sense of havin! outlived their purpose80 ut seem adly desi!ned for
that purpose1 The verterate eye at its est : say0 the eye of a ha%k or a human : is a super
precision instrument0 capale of feats of fine resolution to rival the est that Ieiss or *ikon can
deliver1 #f it %ere not so0 Ieiss and *ikon %ould e %astin! their time producin! hi!h:resolution
ima!es for our eyes to look at1 On the other hand0 )ermann von )elmholtF0 the !reat nineteenth:
century German scientist 7you could call him a physicist0 ut his contriutions to iolo!y and
psycholo!y %ere !reater80 said0 of the eyeA (#f an optician %anted to sell me an instrument %hich had
all these defects0 # should think myself Euite Hustified in lamin! his carelessness in the stron!est
terms0 and !ivin! him ack his instrument1( One reason %hy the eye seems etter than )elmholtF0
the physicist0 Hud!ed it to e is that the rain does an amaFin! Ho of cleanin! the ima!es up
after%ards0 like a sort of ultra:sophisticated0 automatic Photoshop1 &s far as optics are concerned0
the human eye achieves its IeissG*ikon Euality only in the fovea0 the central part of the retina that
%e use for readin!1 When %e scan a scene0 %e move the fovea over different parts0 seein! each one
in the utmost detail and precision0 and the rain(s (Photoshop( fools us into thinkin! %e are seein!
the %hole scene %ith the same precision1 & top:Euality Ieiss or *ikon really does sho% the %hole
scene %ith almost eEual clarity1
So0 %hat the eye lacks in optics the rain makes up for %ith its sophisticated ima!e:
simulatin! soft%are1 But # haven(t yet mentioned the most !larin! example of imperfection in the
optics1 The retina is ack to front1
#ma!ine a latter:day )elmholtF presented y an en!ineer %ith a di!ital camera0 %ith its
screen of tiny photocells0 set up to capture ima!es proHected directly on to the surface of the screen1
That makes !ood sense0 and oviously each photocell has a %ire connectin! it to a computin!
device of some kind %here ima!es are collated1 "akes sense a!ain1 )elmholtF %ouldn(t send it
But no%0 suppose # tell you that the eye(s (photocells( are pointin! ack%ards0 a%ay from the
scene ein! looked at1 The (%ires( connectin! the photocells to the rain run all over the surface of
the retina0 so the li!ht rays have to pass throu!h a carpet of massed %ires efore they hit the
photocells1 That doesn(t make sense : and it !ets even %orse1 One conseEuence of the photocells
pointin! ack%ards is that the %ires that carry their data someho% have to pass throu!h the retina
and ack to the rain1 What they do0 in the verterate eye0 is all conver!e on a particular hole in the
retina0 %here they dive throu!h it1 The hole filled %ith nerves is called the lind spot0 ecause it is
lind0 ut (spot( is too flatterin!0 for it is Euite lar!e0 more like a lind patch0 %hich a!ain doesn(t
actually inconvenience us much ecause of the (automatic Photoshop( soft%are in the rain1 Once
a!ain0 send it ack0 it(s not Hust ad desi!n0 it(s the desi!n of a complete idiot1
H("an eye
Detail o$ '#hotoells' ?ro%s an% ones@
Or is itB #f it %ere0 the eye %ould e terrile at seein!0 and it is not1 #t is actually very !ood1
#t is !ood ecause natural selection0 %orkin! as a s%eeper:up of countless little details0 came alon!
after the i! ori!inal error of installin! the retina ack%ards0 and restored it to a hi!h:Euality
precision instrument1 #t reminds me of the sa!a of the )ule Space Telescope1 6ou(ll rememer
that0 %hen it %as launched in 29950 the )ule %as discovered to possess a maHor fla%1 O%in! to
an undetected fault in the caliration apparatus %hen it %as ein! !round and polished0 the main
mirror %as sli!htly0 ut seriously0 out of shape1 The telescope %as launched into orit0 and then
discovered to e defective1 #n a darin! and resourceful move0 astronauts %ere dispatched to the
telescope0 and they succeeded in fittin! it %ith %hat amounted to spectacles1 The telescope
thereafter %orked very %ell0 and further improvements %ere effected y three more servicin!
missions1 The point # am makin! is that a maHor desi!n fla% : catastrophic lunder0 even : can e
corrected y suseEuent tinkerin!0 %hose in!enuity and intricacy can0 under the ri!ht
circumstances0 perfectly compensate for the initial error1 #n evolution !enerally0 maHor mutations0
even if they cause improvements in !enerally the ri!ht direction0 almost al%ays reEuire a lot of
suseEuent tinkerin! : a s%eepin!:up operation y lots of small mutations that come alon! later and
are favoured y selection ecause they smooth out the rou!h ed!es left y the initial lar!e mutation1
This is %hy humans and ha%ks see so %ell0 despite the lunderin! fla% in the initial desi!n1
)elmholtF a!ainA
,or the eye has every possile defect that can e found in an optical instrument0 and even
some %hich are peculiar to itselfJ ut they are all so counteracted0 that the inexactness of the ima!e
%hich results from their presence very little exceeds0 under ordinary conditions of illumination0 the
limits %hich are set to the delicacy of sensation y the dimensions of the retinal cones1 But as soon
as %e make our oservations under some%hat chan!ed conditions0 %e ecome a%are of the
chromatic aerration0 the asti!matism0 the lind spots0 the venous shado%s0 the imperfect
transparency of the media0 and all the other defects of %hich # have spoken1
$*#*TE--#GE*T 'ES#G*
This pattern of maHor desi!n fla%s0 compensated for y suseEuent tinkerin!0 is exactly
%hat %e should not expect if there really %ere a desi!ner at %ork1 We mi!ht expect unfortunate
mistakes0 as in the spherical aerration of the )ule mirror0 ut %e do not expect ovious
stupidity0 as in the retina ein! installed ack to front1 Blunders of this kind come not from poor
desi!n ut from history1
& favourite example0 ever since it %as pointed out to me y Professor .1 '1 Currey %hen he
tutored me as an under!raduate0 is the recurrent laryn!eal nerve1L #t is a ranch of one of the cranial
nerves0 those nerves that lead directly from the rain rather than from the spinal cord1 One of the
cranial nerves0 the va!us 7the name means (%anderin!( and it is apt80 has various ranches0 t%o of
%hich !o to the heart0 and t%o on each side to the larynx 7voice ox in mammals81 On each side of
the neck0 one of the ranches of the laryn!eal nerve !oes strai!ht to the larynx0 follo%in! a direct
route such as a desi!ner mi!ht have chosen1 The other one !oes to the larynx via an astonishin!
detour1 #t dives ri!ht do%n into the chest0 loops around one of the main arteries leavin! the heart 7a
different artery on the left and ri!ht sides0 ut the principle is the same80 and then heads ack up the
neck to its destination1
#f you think of it as the product of desi!n0 the recurrent laryn!eal nerve is a dis!race1
)elmholtF %ould have had even more cause to send it ack than the eye1 But0 like the eye0 it makes
perfect sense the moment you for!et desi!n and think history instead1 To understand it0 %e need to
!o ack in time to %hen our ancestors %ere fish1 ,ish have a t%o:chamered heart0 unlike our four:
chamered one1 #t pumps lood for%ard throu!h a i! central artery called the ventral aorta1 The
ventral aorta usually !ives off six pairs of ranches0 leadin! off to the six !ills on either side1 The
lood then passes up throu!h the !ills %here it ecomes richly laced %ith oxy!en1 &ove the !ills0
it is collected y six more pairs of lood vessels into another i! vessel runnin! do%n the middle0
called the dorsal aorta0 %hich feeds the rest of the ody1 The six pairs of !ill arteries are evidence of
the se$mented ody plan of the verterates0 %hich is clearer and more ovious in fish than it is in
us1 ,ascinatin!ly0 it is very ovious in human embryos0 %hose (pharyn!eal arches( are clearly
derived from ancestral !ills0 as one can tell y lookin! at their detailed anatomy1 Of course they
don(t function as !ills0 ut five:%eek human emryos can e re!arded as little pink fishes0 %ith !ills1
# can(t help %onderin!0 once a!ain0 %hy %hales and dolphins0 du!on!s and manatees have not re:
evolved functional !ills1 The fact that0 like all mammals0 they have0 in the pharyn!eal arches0 the
emryonic scaffoldin! to !ro% !ills su!!ests that it should not e too difficult to do so1 # don(t kno%
%hy they haven(t0 ut #(m pretty sure there(s a !ood reason0 and someody either kno%s it or kno%s
ho% to research it1
7haryn'eal arhes in h("an e"*ryo
&ll verterates have a se!mented ody plan0 ut in adult mammals as opposed to emryos
this is readily apparent only in the spinal re!ion0 %here the verterae and the ris0 the lood vessels0
muscle locks 7myotomes8 and nerves all follo% a pattern of modular repetition from front to ack1
Every se!ment of the verteral column has t%o i! nerves sproutin! from the spinal cord on either
side0 called the dorsal root and the ventral root1 These nerves mostly do their usiness0 %hatever it
is0 in the vicinity of the verterae from %hich they sprin!0 ut some shoot off do%n the le!s and
some do%n the arms1
The head and neck too0 follo% the same se!mented plan0 ut it is harder to discern0 even in
fish0 ecause the se!ments0 instead of ein! neatly laid out in a fore:and:aft array as they are in the
spinal column0 have ecome all Humled up over evolutionary time1 #t %as one of the triumphs of
nineteenth: and early t%entieth:century comparative anatomy and emryolo!y to discern the
!hostly footprints of se!ments in the head1 ,or example0 the first !ill arch in Ha%less fishes like
lampreys 7and in emryos of Ha%ed verterates8 corresponds to the Ha%s in those verterates that
have them 7that is0 all modern verterates except lampreys and ha!fishes81
#nsects0 too0 and other arthropods such as crustaceans0 as %e sa% in Chapter 250 have a
se!mented ody plan1 &nd it %as a similar triumph to sho% that the insect head contains : a!ain0 all
Humled up : the first six se!ments of %hat0 in their remote ancestors0 %ould have een a train of
modules Hust like the rest of the ody1 #t %as a triumph of late t%entieth:century emryolo!y and
!enetics to sho% that insect se!mentation and verterate se!mentation0 far from ein! independent
of each other as # %as tau!ht0 are actually mediated y parallel sets of !enes0 the so:called hox
!enes0 %hich are reco!niFaly similar in insects and verterates and many other animals0 and that
the !enes are even laid out in the correct serial order in the chromosomesK That is somethin! none
of my teachers %ould have dreamed of %hen # %as an under!raduate learnin!0 entirely separately0
aout insect and verterate se!mentation1 &nimals of different phyla 7for example0 insects and
verterates8 are much more united than %e ever used to think1 &nd that0 too0 is ecause of shared
ancestry1 The hox plan %as already sketched out in the !rand ancestor of all ilaterally symmetrical
animals1 &ll animals are much closer cousins to each other than %e used to think1
To return to the verterate headA the cranial nerves are elieved to e the much:dis!uised
descendants of se!mental nerves0 %hich0 in our primitive ancestors0 constituted the front end of a
train of dorsal roots and ventral roots0 Hust like those %e still have sproutin! from our spinal column1
&nd the maHor lood vessels of our chest are the messed:aout relics and remnants of the once
clearly se!mental lood vessels servin! the !ills1 6ou could say that the mammalian chest has
messed up the se!mental pattern of the ancestral fish !ills0 in the same kind of %ay as0 earlier0 the
fish head messed up the se!mented pattern of even earlier ancestors1
)uman emryos also have lood vessels supplyin! their (!ills(0 %hich are very similar to
those of fish1 There are t%o ventral aortas0 one on each side0 %ith se!mental aortic arches0 one for
each (!ill( on each side0 connectin! to paired dorsal aortas1 "ost of these se!mental lood vessels
have disappeared y the end of emryonic development0 ut it is clearly apparent ho% the adult
pattern is derived from the emryonic : and also from the ancestral : plan1 #f you %ere to look at a
human emryo aout t%enty:six days after conception you %ould see that the lood supply to the
(!ills( stron!ly resemles the se!mental lood supply to the !ills of a fish1 Over the follo%in! %eeks
of !estation the pattern of lood vessels ecomes simplified y sta!es and loses its ori!inal
symmetry0 and y the time the infant is orn its circulatory system has ecome stron!ly left:iased :
Euite unlike the neat symmetry of the fish:like early emryo1
# %on(t !o into the messy details of %hich of our i! chest arteries are the survivors of %hich
of the six numered !ill arteries1 &ll that %e need to kno%0 to understand the history of our
recurrent laryn!eal nerves0 is that in fish the va!us nerve has ranches that supply the last three of
the six !ills0 and it is natural for them0 therefore0 to pass ehind the appropriate !ill arteries1 There is
nothin! (recurrent( aout these ranchesA they seek out their end or!ans0 the !ills0 y the most direct
and lo!ical route1
'urin! the evolution of the mammals0 ho%ever0 the neck stretched 7fish don(t have necks8
and the !ills disappeared0 some of them turnin! into useful thin!s such as the thyroid and
parathyroid !lands0 and the various other its and pieces that comine to form the larynx1 Those
other useful thin!s0 includin! the parts of the larynx0 received their lood supply and their nerve
connections from the evolutionary descendants of the lood vessels and nerves that0 once upon a
time0 served the !ills in orderly seEuence1 &s the ancestors of mammals evolved further and further
a%ay from their fish ancestors0 nerves and lood vessels found themselves pulled and stretched in
puFFlin! directions0 %hich distorted their spatial relations one to another1 The verterate chest and
neck ecame a mess0 unlike the tidily symmetrical0 serial repetitiveness of fish !ills1 &nd the
recurrent laryn!eal nerves ecame more than ordinarily exa!!erated casualties of this distortion1
The picture opposite0 from a 29;< textook y Berry and )allam0 sho%s ho% the laryn!eal
nerve lacks a detour in a shark1 To illustrate the detour in a mammal0 Berry and )allam chose :
%hat more strikin! example could there eB : a !iraffe1
#n a person0 the route taken y the recurrent laryn!eal nerve represents a detour of perhaps
several inches1 But in a !iraffe0 it is eyond a Hoke : many feet eyond : takin! a detour of perhaps
2= feet in a lar!e adultK The day after 'ar%in 'ay 3559 7his 355th irthday8 # %as privile!ed to
spend the %hole day %ith a team of comparative anatomists and veterinary patholo!ists at the Royal
+eterinary Colle!e near -ondon0 dissectin! a youn! !iraffe that had unfortunately died at a Foo1 #t
%as a memorale day0 almost a surreal experience for me1 The operatin! theatre %as literally a
theatre0 %ith a hu!e plate:!lass %all separatin! the (sta!e( from the raked seats %here veterinary
students %ere %atchin! for hours at a time1 &ll day : it must have een ri!ht out of the normal run
of their experience as students : they sat in the darkened theatre and stared throu!h the !lass at the
rilliantly lit scene0 listenin! to the %ords spoken y the dissectin! team0 %ho all %ore throat
microphones0 as did # and the television production cre% filmin! for a future documentary on
Channel ,our1 The !iraffe %as laid out on the lar!e0 an!led dissectin! tale0 %ith one le! held hi!h
in the air y a hook and pulley0 its enormous and affectin!ly vulnerale neck prominently exposed
under ri!ht li!hts1 &ll of us on the !iraffe side of the !lass %all %ere under strict orders to %ear
oran!e overalls and %hite oots0 %hich someho% enhanced the dream:like Euality of the day1
1aryn'eal ner.e in 'ira$$e an% shar&
#t is testimony to the len!th of the detour taken y the recurrent laryn!eal that different
memers of the team of anatomists %orked simultaneously on different stretches of the nerve : the
larynx near the head0 the recurrence itself near the heart0 and all stations et%een : %ithout !ettin!
in each other(s %ay0 and scarcely needin! to communicate %ith each other1 Patiently they teased out
the entire course of the recurrent laryn!eal nerveA a difficult task that had not0 as far as %e kno%0
een achieved since Richard O%en0 the !reat +ictorian anatomist0 did it in 2;4@1 #t %as difficult0
ecause the nerve is very narro%0 even thread:like in its recurrent portion 7# suppose # should have
kno%n that0 ut it came as a surprise0 nevertheless0 %hen # actually sa% it8 and it is easily missed in
the intricate %e of memranes and muscles that surround the %indpipe1 On its do%n%ard Hourney0
the nerve 7at this point it is undled in %ith the lar!er va!us nerve8 passes %ithin inches of the
larynx0 %hich is its final destination1 6et it proceeds do%n the %hole len!th of the neck efore
turnin! round and !oin! all the %ay ack up a!ain1 # %as very impressed %ith the skill of Professors
Graham "itchell and .oy Reidener!0 and the other experts doin! the dissection0 and # found my
respect for Richard O%en 7a itter foe of 'ar%in8 !oin! up1 The creationist O%en0 ho%ever0 failed
to dra% the ovious conclusion1 &ny intelli!ent desi!ner %ould have hived off the laryn!eal nerve
on its %ay do%n0 replacin! a Hourney of many metres y one of a fe% centimetres1
Puite apart from the %aste of resources involved in makin! such a lon! nerve0 # can(t help
%onderin! %hether !iraffe vocaliFations are suHect to a delay0 like a forei!n correspondent talkin!
over a satellite link1 One authority has said0 ('espite possession of a %ell developed larynx and a
!re!arious nature0 the Giraffe is ale to utter only lo% moans or leats1( & !iraffe %ith a stutter is an
endearin! thou!ht0 ut # %on(t pursue it1 The important point is that this %hole story of the detour is
a splendid example of ho% very far livin! creatures are from havin! een %ell desi!ned1 &nd0 for
an evolutionist0 the important Euestion is %hy natural selection does not do as an en!ineer %ouldA
!o ack to the dra%in! oard and reHi! thin!s in a sensile manner1 #t is the same Euestion %e are
meetin! over and over in this chapter0 and # have attempted to ans%er it in various %ays1 The
recurrent laryn!eal lends itself to an ans%er in terms of %hat economists call (mar!inal cost(1
&s the !iraffe(s neck slo%ly len!thened over evolutionary time0 the cost of the detour :
%hether economic cost or cost in terms of (stutterin!( : !radually increased0 %ith the emphasis on
(!radually(1 The mar$inal cost of each millimetre of increase %as sli$ht1 &s the !iraffe(s neck e!an
to approach its present impressive len!th0 the total cost of the detour mi!ht have e!un to approach
the point %here : hypothetically : a mutant individual %ould survive etter if its descendin!
laryn!eal nerve fires hived themselves off from the va!us undle and hopped across the tiny !ap to
the larynx1 But the mutation needed to achieve this (hop across( %ould have to have constituted a
maHor chan!e : upheaval even : in emryonic development1 +ery proaly0 the necessary mutation
%ould never happen to arise any%ay1 Even if it did0 it mi!ht %ell have disadvanta!es : inevitale in
any maHor upheaval durin! the course of a sensitive and delicate process1 &nd even if these
disadvanta!es mi!ht eventually have een out%ei!hed y the advanta!es of ypassin! the detour0
the mar$inal cost of each millimetre of increased detour compared with the existin$ deto"r is
sli!ht1 Even if a (ack to the dra%in! oard( solution %ould e a etter idea if it could e achieved0
the competin! alternative %as Hust a tiny increase over the existin! detour0 and the mar$inal cost of
this tiny increase %ould have een small1 Smaller0 # am conHecturin!0 than the cost of the (maHor
upheaval( reEuired to rin! aout the more ele!ant solution1
Deto(r "a%e *y laryn'eal ner.e in 'ira$$e
&ll that is eside the main point0 %hich is that the recurrent laryn!eal nerve in any mammal
is !ood evidence a!ainst a desi!ner1 &nd in the !iraffe it stretches from !ood to spectacularK That
iFarrely lon! detour do%n the !iraffe(s neck and ack up a!ain is exactly the kind of thin! %e
expect from evolution y natural selection0 and exactly the kind of thin! %e do not expect from any
kind of intelli!ent desi!ner1
Geor!e C1 Williams is one of the most respected of &merican evolutionary iolo!ists 7his
Euiet %isdom and cra!!y features recall one of the most respected of &merican presidents : %ho
happens to have een orn on the same day as Charles 'ar%in and %as also reno%ned for Euiet
%isdom81 Williams called attention to another detour0 similar to that taken y the recurrent laryn!eal
nerve0 ut at the other end of the ody1 The vas deferens is the pipe that carries sperm from the
testis to the penis1 The most direct route is the fictitious one sho%n on the left:hand side of the
dia!ram opposite1 The actual route taken y the vas deferens is sho%n on the ri!ht of the dia!ram1 #t
takes a ridiculous detour around the ureter0 the pipe that carries urine from the kidney to the
ladder1 #f this %ere desi!ned0 noody could seriously deny that the desi!ner had made a ad error1
But0 Hust as %ith the recurrent laryn!eal nerve0 all ecomes clear %hen %e look at evolutionary
history1 The likely ori!inal position of the testes is sho%n in dotted lines1 When0 in the evolution of
mammals0 the testes descended to their present position in the scrotum 7for reasons that are unclear0
ut are often thou!ht to e associated %ith temperature80 the vas deferens unfortunately !ot hooked
the %ron! %ay over the ureter1 Rather than reroute the pipe0 as any sensile en!ineer %ould have
done0 evolution simply kept on len!thenin! it : once a!ain0 the mar!inal cost of each sli!ht increase
in len!th of detour %ould have een small1 6et a!ain0 it is a eautiful example of an initial mistake
compensated for in a post hoc fashion0 rather than ein! properly corrected ack on the dra%in!
oard1 Examples like this must surely undermine the position of those %ho hanker after (intelli!ent
Ro(te o$ .as %e$erens $ro" testis to #enis
The human ody aounds %ith %hat0 in one sense0 %e could call imperfections ut0 in
another sense0 should e seen as inescapale compromises resultin! from our lon! ancestral history
of descent from other kinds of animal1 #mperfections are inevitale %hen (ack to the dra%in!
oard( is not an option : %hen improvements can e achieved only y makin! ad hoc modifications
to %hat is already there1 #ma!ine %hat a mess the Het en!ine %ould e if Sir ,rank Whittle and 'r
)ans von Ohain0 its t%o independent inventors0 had een forced to aide y a rule that saidA (6ou
are not allo%ed to start %ith a clean sheet on your dra%in! oard1 6ou have to start %ith a propeller
en!ine and chan!e it0 one piece at a time0 scre% y scre%0 rivet y rivet0 from the QancestralQ
propeller en!ine into a QdescendantQ Het en!ine1( Even %orse0 all the intermediates have !ot to fly0
and each one in the chain has !ot to e at least a sli!ht improvement on its predecessor1 6ou can see
that the resultin! Het en!ine %ould e urdened %ith all kinds of historical relics and anomalies and
imperfections1 &nd each imperfection %ould e attended y a cumersome accretion of
compensatory od!es and fixes and klud!es0 each one makin! the est of the unfortunate
prohiition a!ainst !oin! ri!ht ack to the dra%in! oard1
The point is made0 ut a closer look at iolo!ical innovation mi!ht dra% a different analo!y
from the propeller en!ine G Het en!ine case1 &n important innovation 7the Het en!ine in our analo!y8
is Euite likely to evolve not from the old or!an that did the same Ho 7the propeller en!ine in this
case8 ut from somethin! completely different0 %hich performed a completely different function1
&s a nice example0 %hen our fish ancestors took to reathin! air0 they didn(t modify their !ills to
make a lun! 7as do some modern air:reathin! fish0 such as the climin! perch Anabas81 #nstead0
they modified a pouch of the !ut1 &nd later0 y the %ay0 the teleosts : %hich means Hust aout any
fish you are likely to meet0 except sharks and their kind : modified the lun! 7%hich had previously
evolved in ancestors that occasionally reathed air8 to ecome yet another vital or!an0 %hich has
nothin! to do %ith reathin!A the s%im ladder1
The s%im ladder is perhaps the maHor key to the teleosts( success0 and it is %ell %orth a
di!ression to explain it1 #t is an internal ladder filled %ith !as0 %hich can e sensitively adHusted to
keep the fish in hydrostatic eEuilirium at any desired depth1 #f you ever played %ith a Cartesian
'iver as a child you(ll reco!niFe the principle0 ut a teleost fish uses an interestin! variant of it1 &
Cartesian 'iver is a little toy %hose usiness part is a tiny upended cup0 containin! a ule of air0
floatin! at eEuilirium in a ottle of %ater1 The numer of molecules of air in the ule is fixed0
ut you can decrease the volume 7and increase the pressure0 follo%in! Boyle(s -a%L8 y pressin!
do%n on the cork in the ottle1 Or you can increase the volume of air 7and decrease the pressure of
the ule8 y raisin! the cork1 The effect is est achieved %ith one of those stout scre% stoppers
they put on cider ottles1 When you lo%er or raise the stopper0 the diver moves do%n or up until it
reaches its ne% point of hydrostatic eEuilirium1 6ou can coax the diver up and do%n the ottle y
sensitive adHustments to the stopper0 and hence to the pressure1
& fish is a Cartesian 'iver %ith a sutle difference1 The s%im ladder is its (ule( and it
%orks in the same %ay0 except that the numer of molecules of !as in the ladder is not fixed1
When the fish %ants to rise to a hi!her level in the %ater it releases molecules of !as from the lood
into the ladder0 therey increasin! the volume1 When it %ants to sink deeper0 it asors molecules
of !as from the ladder into the lood0 therey decreasin! the volume of the ladder1 The s%im
ladder means that a fish doesn(t have to do muscular %ork0 as a shark does0 in order to stay at a
desired depth1 #t is at hydrostatic eEuilirium at %hatever depth it chooses1 The s%im ladder does
that Ho0 therey freein! up the muscles for active propulsion1 Sharks0 y contrast0 have to keep
s%immin! all the time0 other%ise they %ould sink to the ottom0 admittedly slo%ly ecause they
have special lo%:density sustances in their tissues that keep them moderately uoyant1 The s%im
ladder0 then0 is a coopted lun!0 %hich is itself a coopted !ut pouch 7not0 as you mi!ht have
expected0 a coopted !ill chamer81 &nd in some fish0 the s%im ladder itself is yet further coopted
into a hearin! or!an0 a kind of eardrum1 )istory is %ritten all over the ody0 not Hust once ut
repeatedly0 in exuerant palimpsest1
We(ve een land animals for aout >55 million years0 and %e(ve %alked on our hind le!s for
only aout the last 2 per cent of that time1 ,or 99 per cent of our time on land0 %e(ve had a more:or:
less horiFontal ackone and %alked on four le!s1 We don(t kno% for certain %hat selective
advanta!es accrued to the individuals %ho first rose up and %alked on their hind le!s0 and # am
!oin! to leave that matter aside1 .onathan Din!don has %ritten a %hole ook on the Euestion 7(owly
!ri$in8 and # have considered it in some detail in The Ancestor's Tale1 #t may not have seemed like
a maHor chan!e %hen it happened0 ecause other primates such as chimpanFees0 some monkeys and
the enchantin! lemur +erreaux(s sifaka do it from time to time1 )aitually %alkin! only on t%o le!s
as %e do0 ho%ever0 had far:reachin! ramifications all over the ody0 %hich entailed lots of
compensatory adHustments1 #t could e ar!ued that not a sin!le one or muscle0 any%here in the
ody0 %as spared the necessity to chan!e0 in order to reconcile some detail0 ho%ever oscure0
ho%ever out:of:the:%ay0 and ho%ever indirectly or tenuously connected0 %ith the maHor shift in
!ait1 & similar across:the:oard reHi!!in! must attend each and every maHor chan!e in %ay of life0
from %ater to land0 from land to %ater0 into the air0 under!round1 6ou cannot separate out the
ovious chan!es in the ody and treat them in isolation1 To say that there are ramifications of every
chan!e is an understatement1 There are hundreds0 thousands of ramifications0 and ramifications of
ramifications1 *atural selection is forever t%eakin!0 adHustin! the trim0 (tinkerin!( as the !reat
,rench molecular iolo!ist ,rancois .aco put it1
)ere(s another !ood %ay to look at it1 When there(s a maHor shift in the climate0 say an ice
a!e0 you naturally expect natural selection to adHust the animals to it : !ro% a thicker coat of hair0
for example1 But the external climate is not the only sort of (climate( %e have to consider1 Without
any external chan!e at all0 if a maHor ne% mutation arises0 and is favoured y natural selection0 all
the other !enes in the !enome %ill experience it as a chan!e in the internal (!enetic climate(1 *o less
than a shift in the %eather0 it is a chan!e to %hich they have to adHust1 *atural selection has to come
alon! after%ards0 adHustin! to compensate for a maHor chan!e in !enetic (climate(0 exactly as it
%ould if a chan!e had occurred in the external climate1 The initial shift from a four:le!!ed to a t%o:
le!!ed !ait could even have een (internally( !enerated rather than en!endered y a shift in the
external environment1 Either %ay0 it %ould have initiated a complicated cascade of conseEuences0
each one of %hich necessitated compensatory adHustments of (trim(1
($nintelli!ent desi!n( %ould have een a !ood title for this chapter1 #t mi!ht %ell0 indeed0 e
a %orthy anner for a %hole ook on the imperfections of life as a co!ent indicator of the lack of
delierate desi!n0 and more than one author has independently seiFed upon it1 Of these0 ecause #
love the roust irreverence of &ustralian En!lish 7(So %here did #ntelli!ent 'esi!n sprin! from0 like
a oil on a umB(8 # homed in on the deli!htful ook of Royn Williams0 doyen of Sydney science
roadcasters1 &fter complainin! of the a!ony his o%n ack !ives him every mornin!0 in terms that
%ouldn(t come amiss from a %hin!ein! Pom 7don(t !et me %ron!0 # sympathiFe profoundly80
Williams !oes on0 (nearly all acks could make an instant claim on the %arranty0 if there %ere one1
#f MGodN were responsile for ack desi!n0 you(ll have to concede that it %asn(t one of )is est
moments and must have een a deadline rush Ho at the end of the Six 'ays1( The prolem0 of
course0 is that our ancestors %alked for hundreds of millions of years %ith the ackone held more:
or:less horiFontally0 and it doesn(t take kindly to the sudden readHustment imposed y the last fe%
million1 &nd the point0 once a!ain0 is that a real desi!ner of an upri!ht:%alkin! primate %ould have
!one ack to the dra%in! oard and done the Ho properly0 instead of startin! %ith a Euadruped and
Williams next mentions the pouch of that iconic &ustralian animal the koala0 %hich : not a
!reat idea in an animal that spends its time clin!in! to tree trunks : opens do%n%ards0 instead of
up%ards as in a kan!aroo1 Once a!ain0 the reason is a le!acy of history1 Doalas are descended from
a %omat:like ancestor1 Womats are champion di!!ers0
flin!in! !reat pa%s full of soil ack%ards like an excavator di!!in! out a tunnel1 )ad this
ancestor(s pouch pointed for%ards0 its aies %ould have had eyes and teeth permanently filled %ith
!rit1 So ack%ards it %as and0 %hen one day the creature moved up a tree0 perhaps to exploit a fresh
food source0 the (desi!n( came %ith it0 too complicated to chan!e1
&s %ith the recurrent laryn!eal nerve0 it mi!ht theoretically e possile to chan!e the
emryolo!y of the koala to turn its pouch the other %ay up1 But : #(m !uessin! : the emryolo!ical
upheaval attendant on such a maHor chan!e %ould render the intermediates even %orse off than
koalas copin! %ith the existin! state of affairs1
&nother conseEuence of our o%n shift from Euadruped to iped concerns the sinuses0 %hich
!ive such !rief to many of us 7includin! me at the moment of %ritin!8 ecause their draina!e hole is
in the very last place a sensile desi!ner %ould have chosen1 Williams Euotes an &ustralian
collea!ue0 Professor 'erek 'entonAL (The i! maxillary sinuses or cavities are ehind the cheeks on
either side of the face1 They have their draina!e hole in their top0 %hich is not much of an idea in
terms of usin! !ravity to assist draina!e of fluid1( #n a Euadruped0 the (top( is not the top at all ut the
front0 and the position of the draina!e hole makes much more senseA the le!acy of history0 yet a!ain0
is %ritten all over us1
Williams !oes on to Euote another &ustralian collea!ue0 %ho shares the national !ift for
chuckin! a onFer phrase0 on the #chneumonid %asps0 %hose desi!ner0 if there %ere one0 (must have
een a sadistic astard(1 'ar%in0 althou!h he visited &ustralia as a youn! man0 expressed the same
sentiment in staider0 less antipodean termsA (# cannot persuade myself that a eneficent and
omnipotent God %ould have desi!nedly created the #chneumonidae %ith the express intention of
their feedin! %ithin the livin! odies of caterpillars1( The le!endary cruelty of ichneumon %asps
7also the related di!!er %asps and tarantula %asps8 is a leitmotif %hich %ill recur in the final t%o
chapters of the ook1
# find it hard to articulate %hat # am aout to say0 ut it is somethin! that # have een
thinkin! for a %hile0 and it came to a head durin! that memorale day of the dissection of the
!iraffe1 When %e look at animals from the outside0 %e are over%helmin!ly impressed y the
ele!ant illusion of desi!n1 & ro%sin! !iraffe0 a soarin! alatross0 a divin! s%ift0 a s%oopin!
falcon0 a leafy sea dra!on invisile amon! the sea%eed0 a sprintin! cheetah at full stretch after a
s%ervin!0 pronkin! !aFelle : the illusion of desi!n makes so much intuitive sense that it ecomes a
positive effort to put critical thinkin! into !ear and overcome the seductions of naive intuition1
That(s %hen %e look at animals from the outside1 When %e look inside0 the impression is opposite1
&dmittedly0 an impression of ele!ant desi!n is conveyed y simplified dia!rams in textooks0
neatly laid out and colour:coded like an en!ineer(s lueprint1 But the reality that hits you %hen you
see an animal opened up on a dissectin! tale is very different1 # think it %ould e an instructive
exercise to ask an en!ineer to dra% an improved version of0 say0 the arteries leavin! the heart1 #
ima!ine the result %ould e somethin! like the exhaust manifold of a car0 %ith a neat line of pipes
comin! off in orderly array0 instead of the haphaFard mess that %e actually see %hen %e open a real
"y purpose in spendin! a day %ith the anatomists dissectin! a !iraffe %as to study the
recurrent laryn!eal nerve as an example of evolutionary imperfection1 But # soon realiFed that0
%here imperfection is concerned0 the recurrent laryn!eal is Hust the tip of the iceer!1 The fact that it
takes such a lon! detour drives the point home %ith peculiar force1 That is the aspect that %ould
finally provoke a )elmholtF to send it ack1 But the over%helmin! impression you !et from
surveyin! any part of the innards of a lar!e animal is that it is a messK *ot only %ould a desi!ner
never have made a mistake like that nervous detourJ a decent desi!ner %ould never have perpetrated
anythin$ of the shamles that is the criss:crossin! maFe of arteries0 veins0 nerves0 intestines0 %ads
of fat and muscle0 mesenteries and more1 To Euote the &merican iolo!ist Colin Pittendri!h0 the
%hole thin! is nothin! ut a (patch%ork of makeshifts pieced to!ether0 as it %ere0 from %hat %as
availale %hen opportunity knocked0 and accepted in the hindsi!ht0 not the foresi!ht0 of natural
L #n Britain0 (EsE1( meant 7still means0 althou!h the usa!e is rapidly ecomin! extinct8
(!entleman(0 not (la%yer( as 7# recently discovered8 it means in &merica1 # have even encountered
female &merican la%yers referrin! to themselves as (EsE1(1 This seems to En!lish people aout as
odd as &mericans must find the desi!nation of the first female -a% -ord 7British eEuivalent of
Supreme Court .ustice8 as (-ord .ustice EliFaeth Butler:Sloss(1 The En!lish use of (EsE1( seems
even odder to many in the rest of the %orld1 # am told that the (E( pi!eonhole in hotels the %orld
over is replete %ith undelivered letters lookin! for a "r (EsE(1
L The connection %ith the le!endary Sirens may e the hait0 %hich they share %ith their
land relatives the elephants0 of sucklin! their youn! from pectoral reasts1 Perhaps sexually
frustrated sailors %ho had een at sea for a very lon! time %itnessed this from a distance and
mistook them for %omen1 Sirenians have sometimes een lamed for the mermaid le!end1
L -arvae destined to ecome Eueens are fed special elixirs secreted y !lands in the nurse
%orkers( heads1 #t is very important that the difference et%een Eueens and %orkers is
environmentally0 not !enetically0 determined1 # have explained %hy at len!th in The Selfish Gene1
L 6es0 tro!loite0 not tro!lodyte0 %hich means somethin! less extreme1
L This is especially true of mutations of lar!e effect1 Think of a delicate machine0 like a
radio or a computer1 & lar!e mutation is eEuivalent to kickin! it %ith a honailed oot0 or cuttin! a
%ire at random and reconnectin! it in a different place1 #t Hust mi$ht improve its performance0 ut it
is not very likely1 & small mutation0 on the other hand0 is eEuivalent to makin! a tiny adHustment to0
say0 one resistor0 or to the tunin! kno of a radio1 The smaller the mutation0 the more closely the
proaility of improvement approaches =5 per cent1
L #t is also a favourite of my collea!ue .erry Coyne1 Why Eol"tionis Tr"e !ives a
%onderfully clear discussion of this example0 %hich # recommend0 alon! %ith the rest of his
excellent ook1
L Boyle(s -a% states that0 for a fixed Euantity of !as at a !iven temperature0 the pressure is
inversely proportional to the volume1 # have never for!otten Boyle(s -a% since my class at school0
,orm >B20 %as tau!ht a sin!le lesson y the school(s senior science master0 %hose name %as BunHy1
)e %as standin! in for Bufty0 our usual physics teacher0 and %e %ron!ly thou!ht that0 ecause of
BunHy(s extreme a!e 7as %e thou!ht8 and extreme short si!ht 7as %as ovious from his hait of
readin! a ook in contact %ith his nose80 %e could i!nore his discipline and tease him1 )o% %ron!
%e %ere1 )e kept the %hole lot of us in for an extra detention lesson that afternoon0 %hich he e!an
y makin! us %rite in our noteooksA (OHect of the lessonA To teach >B2 !ood manners and Boyle(s
L *ot to e confused %ith another &ustralian0 "ichael 'enton0 eloved of creationists %ho
conveniently overlook the fact that0 in his second ook0 .at"re's Destiny0 he recanted his earlier
anti:evolutionary stance0 %hile remainin! theistic1
C)&PTER 23
&R"S R&CES &*' (E+O-$T#O*&R6 T)EO'#C6(
E 6ES and nerves0 sperm tues0 sinuses and acks are poorly desi!ned from the point of
vie% of individual %elfare0 ut the imperfections make perfect sense in the li!ht of evolution1 The
same applies to the lar!er economy of nature1 &n intelli!ent creator mi!ht e expected to have
desi!ned not Hust the odies of individual animals and plants ut also %hole species0 entire
ecosystems1 *ature mi!ht e expected to e a planned economy0 carefully desi!ned to eliminate
extrava!ance and %aste1 #t isn(t0 and this chapter %ill sho% it1
The natural economy is solar:po%ered1 Photons from the sun rain do%n upon the entire
daytime surface of the planet1 "any photons do nothin! more useful than heat up a rock or a sandy
each1 & fe% find their %ay into an eye : yours0 or mine0 or the compound eye of a shrimp or the
paraolic reflector eye of a scallop1 Some may happen to fall on a solar panel : either a man:made
one like those that0 in a fit of !reen Feal0 # have Hust installed on my roof to heat the ath%ater0 or a
!reen leaf0 %hich is nature(s solar panel1 Plants use solar ener!y to drive (uphill( chemical syntheses0
manufacturin! or!anic fuels0 primarily su!ars1 ($phill( means that the synthesis of su!ar needs
ener!y to drive itJ y the same token0 the su!ar can later e (urned( in a (do%nhill( reaction that
releases 7a fraction of8 the ener!y a!ain to do useful %ork0 for example muscular %ork0 or the %ork
of uildin! a !reat tree trunk1 The (do%nhill( and (uphill( analo!y is %ith %ater flo%in! do%nhill
from a hi!h tank and drivin! %ater %heels to do useful %orkJ or ein! ener!etically pumped uphill
into the hi!h tank0 so that it can later e used to drive %ater %heels %hen it flo%s do%nhill a!ain1
&t every sta!e of the ener!y economy0 %hether uphill or do%nhill0 some ener!y is lost : no ener!y
transaction is ever perfectly efficient1 That is %hy patent offices don(t need even to look at desi!ns
for perpetual motion machinesA they are implacaly and forever impossile1 6ou can(t use the
do%nhill ener!y from a %ater %heel to pump the same amount of %ater uphill a!ain so that it can
drive the %ater %heel1 There must al%ays e some ener!y fed in from outside to compensate for the
%asta!e : and that is %here the sun comes in1 #(ll return to this important theme in Chapter 241
"uch of the land surface of the Earth is covered y !reen leaves0 %hich constitute a many:
layered catchment for photons1 #f a photon is not cau!ht y one leaf0 it has a !ood chance of ein!
cau!ht y the one elo%1 #n a dense forest0 not many photons make it to the !round uncau!ht0
%hich is exactly %hy mature forests are such dark places in %hich to %alk1 "ost of the photons that
constitute our planet(s minute share of the sun(s rays hit %ater0 and the surface layers of the sea
s%arm %ith sin!le:celled !reen plants to catch them1 Whether at sea or on land0 the chemical
process that traps photons and uses them to drive (uphill( ener!y:consumin! chemical reactions0
manufacturin! convenient ener!y:stora!e molecules such as su!ars and starch0 is called
photosynthesis1 #t %as invented0 more than a illion years a!o0 y acteriaJ and !reen acteria still
underlie most photosynthesis1 # can say this ecause the chloroplasts : tiny !reen photosynthetic
en!ines that actually do the usiness of photosynthesis in all leaves : are themselves the direct
descendants of !reen acteria1 #ndeed0 since they still autonomously reproduce themselves after the
manner of acteria0 %ithin plant cells0 %e can Hustly say that they still are acteria0 aleit heavily
dependent on the leaves that house them and to %hich they !ive their colour1 #t appears that
ori!inally free:livin! !reen acteria %ere hiHacked into plant cells0 %here they eventually evolved
into %hat %e no% call chloroplasts1
&nd it is a neatly symmetrical fact that0 Hust as the uphill chemistry of life is mostly taken
care of y !reen acteria thrivin! inside plant cells0 so too the do%nhill chemistry of metaolism :
the slo% urnin! of su!ars and other fuels to release ener!y in cells of oth animals and plants : is
the special expertise of another class of acteria0 once free:livin! ut no% reproducin! themselves
in lar!er cells0 %here they are kno%n as mitochondria1 "itochondria and chloroplasts0 descended
from different kinds of acteria0 each uilt up their complementary chemical %iFardries illions of
years efore the existence of any livin! or!anism visile to the naked eye1 Both %ere later
shan!haied for their chemical skills0 and today they multiply inside the liEuid interiors of the much
lar!er and more complicated cells of creatures i! enou!h for us to see and touch : plant cells in the
case of chloroplasts0 plant and animal cells in the case of mitochondria1
The solar ener!y captured y chloroplasts in plants lies at the ase of complicated food
chains0 in %hich the ener!y passes from plants throu!h herivores0 %hich may e insects0 throu!h
carnivores0 %hich may e insects or insectivores as %ell as %olves and leopards0 throu!h scaven!ers
such as vultures and dun! eetles0 and eventually to a!ents of decay such as fun!i and acteria1 &t
every sta!e of these food chains0 some of the ener!y is %asted as heat as it passes throu!h0 %hile
some of it is used to drive iolo!ical processes such as muscle contraction1 *o ne% ener!y is added
after the initial input from the sun1 With a fe% interestin! ut minor exceptions such as the deniFens
of deep ocean (smokers( %hose ener!y comes from volcanic sources0 all the ener!y that drives life
comes ultimately from sunli!ht0 trapped y plants1
-ook at a sin!le tall tree standin! proud in the middle of an open area1 Why is it so tallB *ot
to e closer to the sunK That lon! trunk could e shortened until the cro%n of the tree %as splayed
out over the !round0 %ith no loss in photons and hu!e savin!s in cost1 So %hy !o to all that expense
of pushin! the cro%n of the tree up to%ards the skyB The ans%er eludes us until %e realiFe that the
natural haitat of such a tree is a forest1 Trees are tall to overtop rival trees : of the same and other
species1 'on(t e misled %hen you see a tree in an open field or !arden that has leafy ranches all
the %ay do%n to the !round1 #t has that %ell:rounded shape so eloved of ser!eant instructors
ecause it is in an open field or !arden1 L 6ou are seein! it out of its natural haitat0 %hich is a
dense forest1 The natural shape of a forest tree is tall and are:trunked0 %ith most of the ranches
and leaves near the top : in the canopy %hich ears the runt of the photon rain1 &nd no%0 here(s an
odd thou!ht1 #f only all the trees in the forest could come to some a!reement : like a trades union
restrictive practice : to !ro% no hi!her than0 say0 25 feet0 every one %ould enefit1 The entire
community : the entire ecosystem : could !ain from the savin!s in %ood0 and ener!y0 %hich are
consumed in uildin! up those to%erin! and costly trunks1
The difficulty of cultivatin! such a!reements of mutual restraint is %ell kno%n0 even in
human affairs %here %e can potentially deploy the !ift of foresi!ht1 & familiar example is a
su!!ested a!reement to sit0 rather than stand0 %hen %atchin! a spectacle such as a horse race1 #f
everyody sat0 tall people %ould still !et a etter vie% than short people0 Hust as they %ould if
everyody stood0 ut %ith the advanta!e that sittin! is more comfortale for everyody1 The
prolems start %hen one short person sittin! ehind a tall person stands0 to !et a etter vie%1
#mmediately0 the person sittin! ehind him stands0 in order to see anythin! at all1 & %ave of
standin! s%eeps around the field0 until everyody is standin!1 #n the end0 everyody is %orse off
than they %ould e if they had all stayed sittin!1
#n a typical mature forest0 the canopy can e thou!ht of as an aerial meado%0 Hust like a
rollin! !rassland prairie0 ut raised on stilts1 The canopy is !atherin! solar ener!y at much the same
rate as a !rassland prairie %ould1 But a sustantial proportion of the ener!y is (%asted( y ein! fed
strai!ht into the stilts0 %hich do nothin! more useful than loft the (meado%( hi!h in the air0 %here it
picks up exactly the same harvest of photons as it %ould : at far lo%er cost : if it %ere laid flat on
the !round1
&nd this rin!s us face to face %ith the difference et%een a desi!ned economy and an
evolutionary economy1 #n a desi!ned economy there %ould e no trees0 or certainly no very tall
treesA no forests0 no canopy1 Trees are a %aste1 Trees are extrava!ant1 Tree trunks are standin!
monuments to futile competition : futile if %e think in terms of a planned economy1 But the natural
economy is not planned1 #ndividual plants compete %ith other plants0 of the same and other species0
and the result is that they !ro% taller and taller0 far taller than any planner %ould recommend1 *ot
indefinitely taller0 ho%ever1 There comes a point %hen !ro%in! another foot taller0 althou!h it
confers a competitive advanta!e0 costs so much that the individual tree doin! it actually ends up
%orse off than its rivals that for!o the extra foot1 #t is the alance of costs and enefits to the
individual trees that finally determines the hei!ht to %hich trees are pressed to !ro%0 not the
enefits that a rational planner could calculate for the trees as a !roup1 &nd of course the alance
ends up at a different maximum in different forests1 The Pacific Coast red%oods 7see them efore
you die8 have proaly never een exceeded1
#ma!ine the fate of a hypothetical forest : let(s call it the ,orest of ,riendship : in %hich0 y
some mysterious concordat0 all the trees have someho% mana!ed to achieve the desirale aim of
lo%erin! the entire canopy to 25 feet1 The canopy looks Hust like any other forest canopy except that
it is only 25 feet hi!h instead of 255 feet1 ,rom the point of vie% of a planned economy0 the ,orest
of ,riendship is more efficient as a forest than the tall forests %ith %hich %e are familiar0 ecause
resources are not put into producin! massive trunks that have no purpose apart from competin! %ith
other trees1
But no%0 suppose one mutant tree %ere to sprin! up in the middle of the ,orest of
,riendship1 This ro!ue tree !ro%s mar!inally taller than the (a!reed( norm of 25 feet1 #mmediately0
this mutant secures a competitive advanta!e1 &dmittedly0 it has to pay the cost of the extra len!th of
trunk1 But it is more than compensated0 as lon$ as allother trees obey the self0denyin$ ordinance0
ecause the extra photons !athered more than pay the extra cost of len!thenin! the trunk1 *atural
selection therefore favours the !enetic tendency to reak out of the self:denyin! ordinance and !ro%
a it taller0 say to 22 feet1 &s the !enerations !o y0 more and more trees reak the emar!o on
hei!ht1 When0 finally0 all the trees in the forest are 22 feet tall0 they are all %orse off than they %ere
eforeA all are payin! the cost of !ro%in! the extra foot1 But they are not !ettin! any extra photons
for their troule1 &nd no% natural selection favours any mutant tendency to !ro% to0 say 23 feet1
&nd so the trees !o on !ettin! taller and taller1 Will this futile clim to%ards the sun ever come to
an endB Why not trees a mile hi!h0 %hy not .ack(s eanstalkB The limit is set at the hei!ht %here the
mar!inal cost of !ro%in! another foot out%ei!hs the !ain in photons from !ro%in! that extra foot1
We are talkin! individual costs and enefits throu!hout this ar!ument1 The forest %ould
look very different if its economy had een desi!ned for the enefit of the forest as a whole1 #n fact0
%hat %e actually see is a forest in %hich each tree species evolved throu!h natural selection
favourin! indiid"al trees that out:competed rival individual trees0 %hether of their o%n or another
species1 Everythin! aout trees is compatile %ith the vie% that they %ere not desi!ned : unless0 of
course0 they %ere desi!ned to supply us %ith timer0 or to deli!ht our eyes and flatter our cameras
in the *e% En!land ,all1 &nd history is not short of those %ho %ould elieve Hust that0 so let(s turn
to a parallel case0 %here the enefits to humanity are harder to alle!eA the arms race et%een hunters
and hunted1
R$**#*G TO ST&6 #* T)E S&"E P-&CE
The five fastest runners amon! mammal species are the cheetah0 the pron!horn 7often called
(antelope( in &merica althou!h it is not closely related to the (true( antelopes of &frica80 the !nu 7or
%ildeeest0 a true antelope althou!h it doesn(t look much like the others80 the lion0 and the
Thomson(s !aFelle 7another true antelope0 %hich really does look like a standard antelope0 a small
one81 *ote that these top:ranked runners are a mixture of hunted and hunters0 and my point is that
this is no accident1
Cheetahs are said to e capale of acceleratin! from 5 to <5 mph in three seconds0 %hich is
ri!ht up there %ith a ,errari0 a Porsche or a Tesla1 -ions0 too0 have formidale acceleration0 even
etter than !aFelles0 %ho have more stamina and the aility to Hink1 Cats !enerally are uilt for
sprintin!0 and sprin!in! on prey taken una%aresJ do!s0 such as the Cape huntin! do! or the %olf0
for endurance0 for %earin! do%n their prey1 GaFelles and other antelopes have to cope %ith oth
types of predator0 and they perhaps have to compromise1 Their acceleration is not Euite so !ood as a
i! cat(s0 ut their endurance is etter1 By Hinkin!0 a Tommy can sometimes thro% a cheetah off its
stride0 therey postponin! matters until the cheetah has !one eyond its maximum acceleration
phase into the exhausted phase0 %here its poor stamina starts to count1 Successful cheetah hunts
usually end soon after they start0 the cheetah relyin! on surprise and acceleration1 $nsuccessful
cheetah hunts also end early0 %ith the cheetah !ivin! up to save ener!y %hen its initial sprint fails1
&ll cheetah hunts0 in other %ords0 are riefK
*ever mind the details of top speeds and accelerations0 stamina and Hinkin!0 surprise and
sustained pursuit1 The salient fact is that the fastest animals include oth those that hunt and those
that are hunted1 *atural selection drives predator species to ecome ever etter at catchin! prey0 and
it simultaneously drives prey species to ecome ever etter at escapin! them1 Predators and prey are
en!a!ed in an evolutionary arms race0 run in evolutionary time1 The result has een a steady
escalation in the Euantity of economic resources that animals0 on oth sides0 spend on the arms race0
at the expense of other departments of their odily economy1 )unters and hunted alike !et steadily
etter eEuipped to outrun 7surprise0 out%it0 etc18 the other side1 But improved eEuipment to outrun
doesn(t oviously translate into improved success in outrunnin! : for the simple reason that the
other side in the arms race is up!radin! its eEuipment tooA that is the hallmark of an arms race1 6ou
could say0 as the Red Pueen said to &lice0 that they have to run as fast as they can Hust to stay in the
same place1
'ar%in %as %ell a%are of evolutionary arms races0 althou!h he didn(t use the phrase1 "y
collea!ue .ohn Dres and # pulished a paper on the suHect in 29@90 in %hich %e attriuted the
phrase (armament race( to the British iolo!ist )u!h Cott1 Perhaps si!nificantly0 Cott pulished his
ook0 Adaptie #oloration in Animals0 in 29>50 in the depths of the Second World WarA
Before assertin! that the deceptive appearance of a !rasshopper or utterfly is unnecessarily
detailed0 %e must first ascertain %hat are the po%ers of perception and discrimination of the insects(
natural enemies1 *ot to do so is like assertin! that the armour of a attle:cruiser is too heavy0 or the
ran!e of her !uns too !reat0 %ithout inEuirin! into the nature and effectiveness of the enemy(s
armament1 The fact is that in the primeval stru!!le of the Hun!le0 as in the refinements of civiliFed
%arfare0L %e see in pro!ress a !reat evolutionary armament race : %hose results0 for defence0 are
manifested in such devices as speed0 alertness0 armour0 spinescence0 urro%in! haits0 nocturnal
haits0 poisonous secretions0 nauseous taste0 and procryptic0 aposematic0 and mimetic colorationJ
and for offence0 in such counter:attriutes as speed0 surprise0 amush0 allurement0 visual acuity0
cla%s0 teeth0 stin!s0 poison fan!s0 and anticryptic and allurin! coloration1 .ust as !reater speed in the
pursued has developed in relation to increased speed in the pursuerJ or defensive armour in relation
to a!!ressive %eaponsJ so the perfection of concealin! devices has evolved in response to increased
po%ers of perception1
*ote that the arms race is run in evolutionary time1 #t is not to e confused %ith the race
et%een an individual cheetah0 say0 and a !aFelle0 %hich is run in real time1 The race in
evolutionary time is a race to uild up eEuipment for races run in real time1 &nd %hat that actually
means is that !enes for makin! the eEuipment to outsmart or outrun the other side uild up in the
!ene pools on the t%o sides1 Second : and this is a point that 'ar%in himself kne% %ell : the
eEuipment for runnin! fast is used to outrun rials of the same species0 %ho are fleein! from the
same predator1 The %ell:kno%n Hoke0 %hich has an almost &esopian rin! to it0 aout the runnin!
shoes and the ear is apposite1 L When a cheetah chases a herd of !aFelles0 it may e more
important for an individual !aFelle to outrun the slo%est memer of the herd than to outrun the
*o% that # have introduced the terminolo!y of the arms race0 you can see that trees in a
forest0 too0 are en!a!ed in one1 #ndividual trees are racin! to%ards the sun0 a!ainst their immediate
nei!hours in the forest1 This race is particularly keen %hen an old tree dies and leaves a vacant slot
in the canopy1 The echoin! crash of an old tree fallin! is the startin! !un for a race0 in real time
7althou!h a slo%er real time than %e animals are accustomed to80 et%een saplin!s that have een
%aitin! for Hust such a chance1 &nd the %inner is likely to e an individual tree that is %ell
eEuipped0 y !enes that prospered throu!h ancestral arms races in evolutionary time0 to !ro% fast
and hi!h1
The arms race et%een species of forest trees is a symmetrical race1 Both sides are tryin! to
achieve the same thin!A a place in the canopy1 The arms race et%een predators and prey is an
asymmetric arms raceA an arms race et%een %eapons of attack and %eapons of defence1 The same
is true of the arms race et%een parasites and hosts1 &nd there are even0 thou!h it may seem
surprisin!0 arms races et%een males and females %ithin a species0 and et%een parents and
One thin! aout arms races that mi!ht %orry enthusiasts for intelli!ent desi!n is the heavy
dose of futility that loads them do%n1 #f %e are !oin! to postulate a desi!ner of the cheetah0 he has
evidently put every ounce of his desi!nin! expertise into the task of perfectin! a superlative killer1
One look at that ma!nificent runnin! machine leaves us in no dout1 The cheetah0 if %e are !oin! to
talk desi!n at all0 is superly desi!ned for killin! !aFelles1 But the very same desi!ner has eEually
evidently strained every nerve to desi!n a !aFelle that is superly eEuipped to escape from those
very same cheetahs1 ,or heaven(s sake0 %hose side is the desi!ner onB When you look at the
cheetah(s taut muscles and flexin! ackone0 you must conclude that the desi!ner %ants the cheetah
to %in the race1 But %hen you look at the sprintin!0 Hinkin!0 dod!in! !aFelle0 you reach exactly the
opposite conclusion1 'oes the desi!ner(s left hand not kno% %hat his ri!ht hand is doin!B #s he a
sadist0 %ho enHoys the spectator sport and is forever uppin! the ante on oth sides to increase the
thrill of the chaseB 'id )e %ho made the lam make theeB
#s it really part of the divine plan that the leopard shall lie do%n %ith the kid0 and the lion eat
stra% like the oxB #n that case0 %hat price the formidale carnassial teeth0 the murderous cla%s of
the lion and the leopardB Whence the reathtakin! speed and a!ile escapolo!y of the antelope and
the FeraB *eedless to say0 no such prolems arise on the evolutionary interpretation of %hat is
!oin! on1 Each side is stru!!lin! to out%it the other ecause0 on oth sides0 those individuals %ho
succeed %ill automatically pass on the !enes that contriuted to their success1 #deas of (futility( and
(%aste( sprin! to our minds ecause %e are human0 and capale of lookin! at the %elfare of the
%hole ecosystem1 *atural selection cares only for the survival and reproduction of individual !enes1
#t(s like the trees in the forest1 .ust as each tree has an economy0 in %hich !oods that are put
into trunks are not availale for fruits or leaves0 so cheetahs and !aFelles each have their o%n
internal economy1 Runnin! fast is costly0 not Hust in ener!y ultimately %run! from the sun ut in the
materials that !o into the makin! of muscles0 ones and sine%s : the machinery of speed and
acceleration1 The food that a !aFelle in!ests in the form of plant material is finite1 Whatever is spent
on muscles and lon! le!s for runnin! has to e taken a%ay from some other department of life0 such
as makin! aies0 on %hich the animal mi!ht ideally (prefer( to spend its resources1 There is an
extremely complicated alance of compromises to e micro:mana!ed1 We can(t kno% all the details
ut %e do kno% 7it is an unreakale la% of economics8 that it is possile to spend too m"ch on one
department of life0 therey takin! resources a%ay from some other department of life1 &n individual
that puts more than the ideal amount into runnin! may save its o%n skin1 But in the 'ar%inian
stakes it %ill e out:competed y a rival individual of the same species0 %ho skimps a little on
runnin! speed and hence incurs a !reater risk of ein! eaten0 ut %ho !ets the alance ri!ht and
ends up %ith more descendants to pass on the !enes for !ettin! the alance ri!ht1
#t isn(t Hust ener!y and costly materials that have to e correctly alanced1 There(s also riskA
and risk0 too0 is no stran!er to the calculations of economists1 -e!s that are lon! and thin are !ood at
runnin! fast1 #nevitaly0 they are also !ood at reakin!1 &ll too re!ularly a racehorse %ill reak a
le! in the heat of a race0 and usually is promptly executed1 &s %e sa% in Chapter 40 the reason they
are so vulnerale is that they have een overred to e fast0 at the expense of everythin! else1
GaFelles and cheetahs have also een selectively red for speed : naturally0 not artificially selected :
and they too %ould e vulnerale to fractures if nature %ere to overreed them for speed1 But nature
never overreeds for anythin!1 *ature !ets the alance ri!ht1 The %orld is full of !enes for !ettin!
the alance ri!htA that is %hy they are thereK What it means in practice is that individuals %ith a
!enetic tendency to develop exceptionally lon! and spindly le!s0 %hich are admittedly superior for
runnin!0 are less likely to pass on their !enes0 on avera!e0 than sli!htly slo%er individuals %hose
less spindly le!s are less likely to reak1 This is Hust one hypothetical example of the many
hundreds of trade:offs and compromises that all animals and plants Hu!!le1 They Hu!!le %ith risks
and they Hu!!le %ith economic trade:offs1 #t is0 of course0 not the individual animals and plants that
do the Hu!!lin! and alancin!1 #t is the relative numers of alternative !enes in !ene pools that are
Hu!!led and alanced0 y natural selection1
&s you %ould expect0 the optimum compromise in a tradeoff is not fixed1 #n !aFelles0 the
trade:off et%een runnin! speed and other demands %ithin the economy of the ody %ill shift its
optimum dependin! upon the prevalence of carnivores in the area1 #t(s the same story as for the
!uppies of Chapter =1 #f there are fe% predators around0 the !aFelle(s optimum le! len!th %ill
shortenA the most successful individuals %ill e the ones %hose !enes predispose them to shunt
some ener!y and material a%ay from le!s and into0 say0 makin! aies0 or layin! do%n fat for the
%inter1 These are also the individuals %ho are less likely to reak their le!s1 Conversely0 if the
numer of predators increases0 the optimum alance %ill shift to%ards lon!er le!s0 !reater dan!er of
fractures0 and less ener!y and material to spend on those aspects of the ody(s economy that are not
concerned %ith runnin! fast1
&nd Hust the same kinds of implicit calculation %ill alance up the optimum compromises in
the predators1 & cheetah %ho reaks her le! %ill undoutedly die of starvation0 and so %ill her cus1
But0 dependin! on ho% difficult it is to find a meal0 the risk of failin! to catch enou!h food if she
runs too slo%ly may out%ei!h the risk of reakin! a le! throu!h ein! eEuipped %ith the
%here%ithal to run too fast1
Predators and prey are locked in an arms race in %hich each side is un%ittin!ly pressin! the
other to shift its optimum : in the economic and risk compromises of life : further and further in the
same directionA either literally in the same direction0 for example to%ards increased runnin! speedJ
or in the same direction in the looser sense of ein! aimed at the predatorGprey arms race rather than
some other department of life such as milk production1 Given that oth sides have to alance the
risks of0 say0 runnin! too fast 7reakin! le!s or skimpin! on the other parts of the odily economy8
a!ainst the risks of runnin! too slo%ly 7failin! to catch prey0 or failin! to escape0 respectively80 each
side is pushin! the other in the same direction0 in a sort of !rim folie a de"x1
Well0 perhaps folie 7madness8 doesn(t Euite do Hustice to the seriousness of the matter0 for the
penalty of failure on either side is death : murder on the side of the prey0 starvation on the side of
the predator1 But a de"x captures handily the feelin! that0 if only hunter and hunted could sit do%n
to!ether and hammer out a sensile a!reement0 everyody %ould e etter off1 .ust as %ith the trees
in the ,orest of ,riendship0 it is easy to see ho% such a compact %ould enefit them0 if only it could
e made to stick1 The same sense of futility as %e encountered in the forest pervades the
predatorGprey arms race1 Over evolutionary time0 predators !et etter at catchin! prey0 %hich
prompts prey animals to !et etter at evadin! capture1 Both sides in parallel improve their
e3"ipment to survive0 ut neither necessarily survives any etter : ecause the other side is
improvin! its eEuipment too1
On the other hand0 it is easy to see ho% a central planner0 %ith the %elfare of the %hole
community at heart0 mi!ht umpire an a!reement in the follo%in! terms0 alon! the lines of the ,orest
of ,riendship1 -et oth sides (a!ree( to scale do%n their armouryA oth sides shift resources to other
departments of life0 and all %ill do etter as a result1 .ust the same0 of course0 can happen in a
human arms race1 We %ouldn(t need our fi!hters if you didn(t have your omers1 6ou %ouldn(t
need your missiles if %e didn(t have ours1 We could oth save illions if %e halved our armaments
spendin! and put the money into plou!hshares1 &nd no%0 havin! halved our arms ud!et and
reached a stale stand:off0 let(s halve it a!ain1 The trick is to do it in synchrony %ith each other0 so
that each side remains exactly as %ell eEuipped to counter the other(s steadily de:escalatin! arms
ud!et1 Such planned de:escalation has to e Hust that : planned1 &nd0 once a!ain0 planned is
precisely %hat evolution is not1 .ust as %ith the trees in the forest0 escalation is inevitale0 ri!ht up
until the moment %hen it no lon!er pays a typical individual to escalate any further1 Evolution0
unlike a desi!ner0 never stops to consider %hether there mi!ht e a etter %ay : a mutualistic %ay :
for all concerned0 rather than ilateral escalation for a selfish advanta!eA an advanta!e that is
neutraliFed precisely ecause the escalation is mutual1
The temptation to think like a planner has lon! een rife amon! (pop ecolo!ists(0 and even
academic ecolo!ists sometimes come perilously close to it1 The temptin! notion of (prudent
predators(0 for example0 %as dreamed up not y some tree:hu!!in! airhead ut y a distin!uished
&merican ecolo!ist1
The idea of prudent predators is this1 Everyody kno%s that0 from the point of vie% of
humanity as a %hole0 %e(d e etter off if %e all refrained from overfishin! an important food
species0 such as the cod0 to extinction1 That is %hy !overnments and *GOs in stately conclave meet
to dra% up Euotas and restrictions1 That is %hy the precise mesh siFe of fishin! nets is minutely
specified y !overnment decree0 and that is %hy !unoats patrol the seas in pursuit of dissentin!
tra%lermen1 We humans0 on our !ood days and %hen properly policed0 are (prudent predators(1
Therefore : or so it seems to certain ecolo!ists : shouldn(t %e expect %ild predators0 like %olves or
lions0 to e prudent predators tooB The ans%er is no1 *o1 *o1 *o1 &nd it is %orth%hile
understandin! %hy0 ecause it(s an interestin! point0 one that the forest trees and this %hole chapter
should have prepared us for1
& planner : an ecosystem desi!ner %ith the %elfare of the %hole community of %ild animals
at heart : could indeed calculate an optimum cullin! policy0 %hich lions0 for example0 should ideally
adopt1 'on(t take more than a certain Euota from any one species of antelope1 Spare pre!nant
females0 and don(t take youn! adults full of reproductive potential1 &void eatin! memers of rare
species0 %hich mi!ht e in dan!er of extinction and mi!ht come in useful in future0 if conditions
chan!e1 #f only all the lions in the country %ould aide y the a!reed norms and Euotas0 carefully
calculated to e (sustainale(0 %ouldn(t that e niceB &nd so sensile1 #f onlyK
Well0 it %ould e sensile0 and it is %hat a desi!ner %ould prescrie0 at least if he had the
%elfare of the ecosystem as a %hole at heart1 But it isn(t %hat natural selection %ould prescrie
7mainly ecause natural selection0 lackin! foresi!ht0 cannot prescribe at all8 and it isn(t %hat
happensK )ere(s %hy0 and it is a!ain the same story as for the trees in the forest1 #ma!ine that0 y
some Euirk of leonine diplomacy0 a maHority of lions in an area someho% mana!ed to a!ree to limit
their huntin! to sustainale levels1 But no%0 suppose that in this other%ise restrained and pulic:
spirited population0 a mutant !ene arose that caused an individual lion to reak a%ay from the
a!reement and exploit the prey population to the uttermost0 even at the risk of drivin! the prey
species extinct1 Would natural selection penaliFe the reellious selfish !eneB &las0 it %ould not1
Offsprin! of the reel lion0 possessors of the reel !ene0 %ould out:compete and out:reproduce their
rivals in the lion population1 Within a fe% !enerations0 the reel !ene %ould spread throu!h the
population and nothin! %ould e left of the ori!inal amicale compact1 )e L %ho !ets the lion(s
share passes on the !enes for doin! so1
But0 the plannin! enthusiast %ill protest0 %hen all the lions are ehavin! selfishly and over:
huntin! the prey species to the point of extinction0 eerybody is %orse off0 even the individual lions
that are the most successful hunters1 $ltimately0 if all the prey !o extinct0 the entire lion population
%ill too1 Surely0 the planner insists0 natural selection %ill step in to stop that happenin!B Once a!ain
alas0 and once a!ain no1 The prolem is that natural selection doesn(t (step in(0 natural selection
doesn(t look into the future0
and natural selection doesn(t choose et%een rival !roups1 #f it did0
there %ould e some chance that prudent predation could e favoured1 *atural selection0 as 'ar%in
realiFed much more clearly than many of his successors0 chooses et%een rival individuals %ithin a
population1 Even if the entire population is divin! to extinction0 driven do%n y individual
competition0 natural selection %ill still favour the most competitive individuals0 ri!ht up to the
moment %hen the last one dies1 *atural selection can drive a population to extinction0 %hile
constantly favourin!0 to the itter end0 those competitive !enes that are destined to e the last to !o
extinct1 The hypothetical planner that # have ima!ined is a certain kind of economist0 a %elfare
economist calculatin! an optimum strate!y for a %hole population0 or an entire ecosystem1 #f %e
must make economic analo!ies0 %e should think instead of &dam Smith(s (invisile hand(1
E+O-$T#O*&R6 T)EO'#C6B
But no% # %ant to leave economics alto!ether1 We shall stay %ith the idea of a planner0 a
desi!ner0 ut our planner %ill e a moral philosopher rather than an economist1 & eneficent
desi!ner mi!ht : you(d idealistically think : seek to minimiFe sufferin!1 This is not incompatile
%ith economic %elfare0 ut the system created %ill differ in detail1 &nd0 once a!ain0 it unfortunately
doesn(t happen in nature1 Why should itB Terrile ut true0 the sufferin! amon! %ild animals is so
appallin! that sensitive souls %ould est not contemplate it1 'ar%in kne% %hereof he spoke %hen
he said0 in a letter to his friend )ooker0 (What a ook a devil(s chaplain mi!ht %rite on the clumsy0
%asteful0 lunderin! lo% and horridly cruel %orks of nature1( The memorale phrase (devil(s
chaplain( !ave me my title for one of my previous ooks0 and in another # put it like thisA
M*Nature is neither kind nor unkind1 She is neither a!ainst sufferin!0 nor for it1 *ature is not
interested in sufferin! one %ay or the other unless it affects the survival of '*&1 #t is easy to
ima!ine a !ene that0 say0 tranEuillises !aFelles %hen they are aout to suffer a killin! ite1 Would
such a !ene e favoured y natural selectionB *ot unless the act of tranEuillisin! a !aFelle improved
that !ene(s chances of ein! propa!ated into future !enerations1 #t is hard to see %hy this should e
so and %e may therefore !uess that !aFelles suffer horrile pain and fear %hen they are pursued to
the death : as most of them eventually are1 The total amount of sufferin! per year in the natural
%orld is eyond all decent contemplation1 'urin! the minute that it takes me to compose this
sentence0 thousands of animals are ein! eaten alive0 others are runnin! for their lives0 %himperin!
%ith fear0 others are ein! slo%ly devoured from %ithin y raspin! parasites0 thousands of all kinds
are dyin! of starvation0 thirst and disease1 #t must e so1 #f there is ever a time of plenty0 this very
fact %ill automatically lead to an increase in population until the natural state of starvation and
misery is restored1
Parasites proaly cause even more sufferin! than predators0 and understandin! their
evolutionary rationale adds to0 rather than miti!ates0 the sense of futility %e experience %hen %e
contemplate it1 # fulminate a!ainst it every time # !et a cold 7# have one no%0 as it happens81 "aye
it is only a minor inconvenience0 ut it is so pointlessK &t least if you are eaten y an anaconda you
can feel that you have contriuted to the %ell:ein! of one of the lords of life1 When you are eaten
y a ti!er0 perhaps your last thou!ht could e0 What immortal hand or eye could frame thy fearful
symmetryB 7#n %hat distant deeps or skies0 urnt the fire of thine eyesB8 But a virusK & virus has
pointless futility %ritten into its very '*& : actually0 R*& in the case of the common cold virus0
ut the principle is the same1 & virus exists for the sole purpose of makin! more viruses1 Well0 the
same is ultimately true of ti!ers and snakes0 ut there it doesn(t seem so futile1 The ti!er and the
snake may e '*&:replicatin! machines ut they are eautiful0 ele!ant0 complicated0 expensive
'*&:replicatin! machines1 #(ve !iven money to preserve the ti!er0 ut %ho %ould think of !ivin!
money to preserve the common coldB #t(s the futility of it that !ets to me0 as # lo% my nose yet
a!ain and !asp for reath1
,utilityB What nonsense1 Sentimental0 human nonsense1 *atural selection is all futile1 #t is
all aout the survival of self:replicatin! instructions for self:replication1 #f a variant of '*&
survives throu!h an anaconda s%allo%in! me %hole0 or a variant of R*& survives y makin! me
sneeFe0 then that is all %e need y %ay of explanation1 +iruses and ti!ers are oth uilt y coded
instructions %hose ultimate messa!e is0 like a computer virus0 ('uplicate me1( #n the case of the cold
virus0 the instruction is executed rather directly1 & ti!er(s '*& is also a (duplicate me( pro!ram0 ut
it contains an almost fantastically lar!e di!ression as an essential part of the efficient execution of
its fundamental messa!e1 That di!ression is a ti!er0 complete %ith fan!s0 cla%s0 runnin! muscles0
stalkin! and pouncin! instincts1 The ti!er(s '*& says0 ('uplicate me y the round:aout route of
uildin! a ti!er first1( &t the same time0 antelope '*& says0 ('uplicate me y the round:aout route
of uildin! an antelope first0 complete %ith lon! le!s and fast muscles0 complete %ith timorous
instincts and finely honed sense or!ans tuned to the dan!er from ti!ers1( Sufferin! is a yproduct of
evolution y natural selection0 an inevitale conseEuence that may %orry us in our more
sympathetic moments ut cannot e expected to %orry a ti!er : even if a ti!er can e said to %orry
aout anythin! at all : and certainly cannot e expected to %orry its !enes1
Theolo!ians %orry aout the prolems of sufferin! and evil0 to the extent that they have
even invented a name0 (theodicy( 7literally0 (Hustice of God(80 for the enterprise of tryin! to reconcile
it %ith the presumed eneficence of God1 Evolutionary iolo!ists see no prolem0 ecause evil and
sufferin! don(t count for anythin!0 one %ay or the other0 in the calculus of !ene survival1
*evertheless0 %e do need to consider the prolem of pain1 Where0 on the evolutionary vie%0 does it
come fromB
Pain0 like everythin! else aout life0 %e presume0 is a 'ar%inian device0 %hich functions to
improve the sufferer(s survival1 Brains are uilt %ith a rule of thum such as0 (#f you experience the
sensation of pain0 stop %hatever you are doin! and don(t do it a!ain1( #t remains a matter for
interestin! discussion %hy it has to e so damned painful1 Theoretically0 you(d think0 the eEuivalent
of a little red fla! could painlessly e raised some%here in the rain0 %henever the animal does
somethin! that dama!es itA picks up a red:hot cinder0 perhaps1 &n imperative admonition0 ('on(t do
that a!ainK( or a painless chan!e in the %irin! dia!ram of the rain such that0 as a matter of fact0 the
animal doesn't do it a!ain0 %ould seem0 on the face of it0 enou!h1 Why the searin! a!ony0 an a!ony
that can last for days0 and from %hich the memory may never shake itself freeB Perhaps !rapplin!
%ith this Euestion is evolutionary theory(s o%n version of theodicy1 Why so painfulB What(s %ron!
%ith the little red fla!B
# don(t have a decisive ans%er1 One intri!uin! possiility is this1 What if the rain is suHect
to opposin! desires and impulses0 and there is some kind of internal tussle et%een themB
SuHectively0 %e kno% the feelin! %ell1 We may e in a conflict et%een0 say0 hun!er and a desire
to e slim1 Or %e may e in a conflict et%een an!er and fear1 Or et%een sexual desire and a shy
fear of reHection0 or a conscience that ur!es fidelity1 We can literally feel the tu! of %ar %ithin us0 as
our conflictin! desires attle it out1 *o%0 ack to pain and its possile superiority over a (red fla!(1
.ust as the desire to e slim can over:rule hun!er0 it is clearly possile to over:rule the desire to
escape pain1 Torture victims may succum eventually0 ut they often !o throu!h a phase of
endurin! considerale pain rather than0 say0 etray their comrades or their country or their ideolo!y1
#n so far as natural selection can e said to (%ant( anythin!0 natural selection doesn(t %ant
individuals to sacrifice themselves for the love of a country0 or for the sake of an ideolo!y or a party
or a !roup or a species1 *atural selection is (a!ainst( individuals over:rulin! the %arnin! sensations
of pain1 *atural selection (%ants( us to survive0 or more specifically0 to reproduce0 and e lo%ed to
country0 ideolo!y or their non:human eEuivalents1 &s far as natural selection is concerned0 little red
fla!s %ill e favoured only if they are never over:ruled1
*o%0 despite philosophical difficulties0 # think that instances %here pain %as over:ruled for
non:'ar%inian reasons : reasons of loyalty to country0 ideolo!y0 etc1 : %ould e more freEuent if
%e had a (red fla!( in the rain rather than real0 full:on0 intolerale pain1 Suppose !enetic mutants
arose %ho could not feel the excruciatin! a!ony of pain ut relied upon a (red fla!( system to keep
them a%ay from odily dama!e1 #t %ould e so easy for them to resist torture0 they(d promptly e
recruited as spies1 Except that it %ould e so easy to recruit a!ents prepared to ear torture that
torture %ould simply stop ein! used as a method of extortion1 But0 in a %ild state0 %ould such
pain:free0 red:fla! mutants survive etter than rival individuals %hose rains do pain in earnestB
Would they survive to pass on the !enes for red:fla! pain sustitutesB Even settin! aside the special
circumstance of torture0 and the special circumstances of loyalty to ideolo!ies0 # think %e can see
that the ans%er mi!ht e no1 &nd %e can ima!ine non:human eEuivalents1
&s a matter of interest0 there are aerrant individuals %ho cannot feel pain0 and they usually
come to a ad end1 (Con!enital insensitivity to pain %ith anhidrosis( 7C#P&8 is a rare !enetic
anormality in %hich the patient lacks pain receptor cells in the skin 7and also : that(s the
(anhidrosis( : doesn(t s%eat81 &dmittedly0 C#P& patients don(t have a uilt:in (red fla!( system to
compensate for the reakdo%n of the pain system0 ut you(d think they could e tau!ht to e
co!nitively a%are of the need to avoid odily dama!e : a learned red fla! system1 &t all events0
C#P& patients succum to a variety of unpleasant conseEuences of their inaility to feel pain0
includin! urns0 reaka!es0 multiple scars0 infections0 untreated appendicitis and scratches to the
eyealls1 "ore unexpectedly0 they also suffer serious dama!e to their Hoints ecause0 unlike the rest
of us0 they don(t shift their posture %hen they have een sittin! or lyin! in one position for a lon!
time1 Some patients set timers to remind themselves to chan!e position freEuently durin! the day1
Even if a (red fla!( system in the rain could e made effective0 there seems to e no reason
%hy natural selection %ould positively favour it over a real pain system Hust ecause it is less
unpleasant1 $nlike our hypothetically eneficent desi!ner0 natural selection is indifferent to the
intensity of sufferin! : except in so far as it affects survival and reproduction1 &nd0 Hust as %e
should expect if the survival of the fittest0 rather than desi!n0 underlies the %orld of nature0 the
%orld of nature seems to take no steps at all to reduce the sum total of sufferin!1 Stephen .ay Gould
reflected on such matters in a nice essay on (*onmoral nature(1 # learned from it that 'ar%in(s
famous revulsion at the #chneumonidae0 %hich # Euoted at the end of the previous chapter0 %as far
from uniEue amon! +ictorian thinkers1
#chneumon %asps0 %ith their hait of paralysin! ut not killin! their victim0 efore layin! an
e!! in it %ith the promise of a larva !na%in! it hollo% from %ithin0 and the cruelty of nature
!enerally0 %ere maHor preoccupations of +ictorian theodicy1 #t(s easy to see %hy1 The female %asps
lay their e!!s in live insect prey0 such as caterpillars0 ut not efore carefully seekin! out %ith their
stin! each nerve !an!lion in turn0 in such a %ay that the prey is paralysed0 ut still stays alive1 #t
must e kept alive to provide fresh meat for the !ro%in! %asp larva feedin! inside1 &nd the larva0
for its part0 takes care to eat the internal or!ans in a Hudicious order1 #t e!ins y takin! out the fat
odies and di!estive or!ans0 leavin! the vital heart and nervous system till last : they are necessary0
you see0 to keep the caterpillar alive1 &s 'ar%in so poi!nantly %ondered0 %hat kind of eneficent
desi!ner %ould have dreamed that upB # don(t kno% %hether caterpillars can feel pain1 # devoutly
hope not1 But %hat # do kno% is that natural selection %ould in any case take no steps to dull their
pain0 if the Ho could e accomplished more economically y simply paralysin! their movements1
Gould Euotes the Reverend William Buckland0 a leadin! nineteenth:century !eolo!ist0 %ho
found consolation in the optimistic spin that he mana!ed to confer on the sufferin! caused y
The appointment of death y the a!ency of carnivora0 as the ordinary termination of animal
existence0 appears therefore in its main results to e a dispensation of enevolenceJ it deducts much
from the a!!re!ate amount of the pain of universal deathJ it arid!es0 and almost annihilates0
throu!hout the rute creation0 the misery of disease0 and accidental inHuries0 and lin!erin! decayJ
and imposes such salutary restraint upon excessive increase of numers0 that the supply of food
maintains perpetually a due ratio to the demand1 The result is0 that the surface of the land and depths
of the %aters are ever cro%ded %ith myriads of animated ein!s0 the pleasures of %hose life are
coextensive %ith its durationJ and %hich throu!hout the little day of existence that is allotted to
them0 fulfill %ith Hoy the functions for %hich they %ere created1
Well0 isn(t that nice for themK
L (#n the army0 %e has three kinds of treesA fir0 poplar0 and ushy top1(
L &n oxymoron if ever there %as one1
L T%o hikers are pursued y a ear1 One hiker runs a%ay0 the other stops to put on his
runnin! shoes1 (&re you madB Even %ith runnin! shoes0 you can(t outrun a !riFFly1( (*o0 ut # can
outrun you1(
L Or she1 The particular case of lions is complicated y the fact that females do most of the
huntin!0 ut males tend to !et (the lion(s share( in any case1 'on(t !et hun! up on (lions( in my
hypothetical example1 Think of a !eneraliFed predator species0 and ima!ine (prudent( individuals
%ho refrain from over:huntin!0 and (imprudent( individuals %ho reak a%ay from the a!reement1
O -oose talk aout 'ar%inian adaptation freEuently founders on the fallacious assumption
7not made explicit0 and the more pernicious in conseEuence8 that evolution has foresi!ht1 Sydney
Brenner0 hero of the #aenorhabditis section of Chapter ;0 has a sardonic %it to match his scientific
rilliance1 # once heard him lampoon the (evolutionary foresi!ht( fallacy y ima!inin! a species in
the Camrian that retained in its !ene pool an other%ise useless protein0 ecause (#t mi!ht come in
handy in the Cretaceous1(
C)&PTER 24
T)ERE #S GR&*'E$R #* T)#S +#EW O, -#,E
$ *-#DE his evolutionist !randfather Erasmus0 %hose scientific verse %as 7some%hat
surprisin!ly0 # have to say8 admired y Words%orth and Colerid!e0 Charles 'ar%in %as not kno%n
as a poet0 ut he produced a lyrical crescendo in the last para!raph of !n the !ri$in of Species1
Thus0 from the %ar of nature0 from famine and death0L the most exalted oHect %hich %e are
capale of conceivin!0 namely0 the production of the hi!her animals0 directly follo%s1 There is
!randeur in this vie% of life0 %ith its several po%ers0 havin! een ori!inally reathed into a fe%
forms or into oneJ and that0 %hilst this planet has !one cyclin! on accordin! to the fixed la% of
!ravity0 from so simple a e!innin! endless forms most eautiful and most %onderful have een0
and are ein!0 evolved1
There(s a lot packed into this famous peroration0 and # %ant to si!n off y takin! it line y
(,RO" T)E W&R O, *&T$RE0 ,RO" ,&"#*E &*' 'E&T)(
Clear:headed as ever0 'ar%in reco!niFed the moral paradox at the heart of his !reat theory1
)e didn(t mince %ords : ut he offered the miti!atin! reflection that nature has no evil intentions1
Thin!s simply follo% from (la%s actin! all around us(0 to Euote an earlier sentence from the same
para!raph1 )e had said somethin! similar at the end of Chapter @ of The !ri$inA
it may not e a lo!ical deduction0 ut to my ima!ination it is far more satisfactory to look at
such instincts as the youn! cuckoo eHectin! its foster:rothers0 : ants makin! slaves0 : the larvae of
ichneumonidae feedin! %ithin the live odies of caterpillars0 : not as specially endo%ed or created
instincts0 ut as small conseEuences of one !eneral la%0 leadin! to the advancement of all or!anic
ein!s0 namely0 multiply0 vary0 let the stron!est live and the %eakest die1
#(ve already mentioned 'ar%in(s revulsion : %idely shared y his contemporaries : in the
face of the female ichneumon %asp(s hait of stin!in! its victim to paralyse ut not kill it0 therey
keepin! the meat fresh for its larva as it eats the live prey from %ithin1 'ar%in0 you(ll rememer0
couldn(t persuade himself that a eneficent creator %ould conceive such a hait1 But %ith natural
selection in the drivin! seat0 all ecomes clear0 understandale and sensile1 *atural selection cares
nau!ht for any comfort1 Why should itB ,or somethin! to happen in nature0 the only reEuirement is
that the same happenin! in ancestral times assisted the survival of the !enes promotin! it1 Gene
survival is a sufficient explanation for the cruelty of %asps and the callous indifference of all natureA
sufficient : and satisfyin! to the intellect if not to human compassion1
6es0 there is !randeur in this vie% of life0 and even a kind of !randeur in nature(s serene
indifference to the sufferin! that inexoraly follo%s in the %ake of its !uidin! principle0 survival of
the fittest1 Theolo!ians may here %ince at this echo of a familiar ploy in theodicy0 in %hich
sufferin! is seen as an inevitale correlate of free %ill1 Biolo!ists0 for their part0 %ill find
(inexoraly( y no means too stron! %hen they reflect : perhaps alon! the lines of my (red fla!(
meditation of the previous chapter : on the iolo!ical function of the capacity to suffer1 #f animals
aren(t sufferin!0 someody isn(t %orkin! hard enou!h at the usiness of !ene survival1
Scientists are human0 and they are as entitled as anyone to revile cruelty and ahor sufferin!1
But !ood scientists like 'ar%in reco!niFe that truths aout the real %orld0 ho%ever distasteful0 have
to e faced1 "oreover0 if %e are !oin! to admit suHective considerations0 there is a fascination in
the leak lo!ic that pervades all of life0 includin! %asps homin! in on the nerve !an!lia do%n the
len!th of their prey0 cuckoos eHectin! their foster rothers 7(Tho% mortherer of the heysu!!e on y
raunche(80 slave:makin! ants0 and the sin!le:minded : or rather Fero:minded : indifference to
sufferin! sho%n y all parasites and predators1 'ar%in %as endin! over ack%ards to console
%hen he concluded his chapter on the stru!!le for survival %ith these %ordsA
&ll that %e can do0 is to keep steadily in mind that each or!anic ein! is strivin! to increase
at a !eometrical ratioJ that each at some period of its life0 durin! some season of the year0 durin!
each !eneration or at intervals0 has to stru!!le for life0 and to suffer !reat destruction1 When %e
reflect on this stru!!le0 %e may console ourselves %ith the full elief0 that the %ar of nature is not
incessant0 that no fear is felt0L that death is !enerally prompt0 and that the vi!orous0 the healthy0 and
the happy survive and multiply1
Shootin! the messen!er is one of humanity(s sillier foiles0 and it underlies a !ood slice of
the opposition to evolution that # mentioned in the #ntroduction1 (Teach children that they are
animals0 and they(ll ehave like animals1( Even if it %ere true that evolution0 or the teachin! of
evolution0 encoura!ed immorality0 that %ould not imply that the theory of evolution %as false1 #t is
Euite astonishin! ho% many people cannot !rasp this simple point of lo!ic1 The fallacy is so
common it even has a name0 the ar$"ment"m ad conse3"entiam : C is true 7or false8 ecause of
ho% much # like 7or dislike8 its conseEuences1
(T)E "OST EC&-TE' OB.ECT W)#C) WE &RE C&P&B-E O, CO*CE#+#*G(
#s (the production of the hi!her animals( really (the most exalted oHect %hich %e are capale
of conceivin!(B %ost exaltedB ReallyB &re there not more exalted oHectsB &rtB SpiritualityB Romeo
and -"lietB General RelativityB The Choral SymphonyB The Sistine ChapelB -oveB
6ou have to rememer that0 for all his personal modesty0 'ar%in nursed hi!h amitions1 On
his %orld:vie%0 everythin! aout the human mind0 all our emotions and spiritual pretensions0 all
arts and mathematics0 philosophy and music0 all feats of intellect and of spirit0 are themselves
productions of the same process that delivered the hi!her animals1 #t is not Hust that %ithout evolved
rains spirituality and music %ould e impossile1 "ore pointedly0 rains %ere naturally selected to
increase in capacity and po%er for utilitarian reasons0 until those hi!her faculties of intellect and
spirit emer!ed as a y:product0 and lossomed in the cultural environment provided y !roup livin!
and lan!ua!e1 The 'ar%inian %orld:vie% does not deni!rate the hi!her human faculties0 does not
(reduce( them to a plane of indi!nity1 #t doesn(t even claim to explain them at the sort of level that
%ill seem particularly satisfyin!0 in the %ay that0 say0 the 'ar%inian explanation of a snake:
mimickin! caterpillar is satisfyin!1 #t does0 ho%ever0 claim to have %iped out the impenetrale : not
even %orth tryin! to penetrate : mystery that must have do!!ed all pre:'ar%inian efforts to
understand life1
But 'ar%in doesn(t need any defence from me0 and #(ll pass over the Euestion of %hether the
production of the hi!her animals is the most exalted oHect %e can conceive0 or merely a very
exalted oHect1 What0 ho%ever0 of the predicateB 'oes the production of the hi!her animals (directly
follo%( from the %ar of nature0 from famine and deathB Well0 yes0 it does1 #t directly follo%s if you
understand 'ar%in(s reasonin!0 ut noody understood it until the nineteenth century1 &nd many
still don(t understand it0 or perhaps are reluctant to do so1 #t is not hard to see %hy1 When you think
aout it0 our o%n existence0 to!ether %ith its post:'ar%inian explicaility0 is a candidate for the
most astonishin! fact that any of us are called upon to contemplate0 in our %hole life0 ever1 #(ll come
to that shortly1
()&+#*G BEE* OR#G#*&--6 BRE&T)E'(
# have lost count of the irate letters # have received from readers of a previous ook0 takin!
me to task for0 as the %riters think0 delierately omittin! the vital phrase0 (y the Creator( after
(reathed(B &m # not %antonly distortin! 'ar%in(s intentionB These Fealous correspondents for!et
that 'ar%in(s !reat ook %ent throu!h six editions1 #n the first edition0 the sentence is as # have
%ritten it here1 Presumaly o%in! to pressure from the reli!ious loy0 'ar%in inserted (y the
Creator( in the second and all suseEuent editions1 $nless there is a very !ood reason to the
contrary0 %hen Euotin! !n the!ri$in of Species # al%ays Euote the first edition1 This is partly
ecause my o%n copy of that historic print run of 203=5 is one of my most precious possessions0
!iven me y my enefactor and friend Charles Simonyi1 But it is also ecause the first edition is the
most historically important1 #t is the one that thumped the +ictorian solar plexus and drove out the
%ind of centuries1 "oreover0 later editions0 especially the sixth0 pandered to more than pulic
opinion1 #n an attempt to respond to various learned ut mis!uided critics of the first edition0
'ar%in acktracked and even reversed his position on a numer of important points that he had
actually !ot ri!ht in the first place1 So0 (havin! een ori!inally reathed( it is0 %ith no mention of
any Creator1
#t seems that 'ar%in re!retted this sop to reli!ious opinion1 #n a letter of 2;<4 to his friend
the otanist .oseph )ooker0 he said0 (But # have lon! re!retted that # truckled to pulic opinion0 and
used the Pentateuchal term of creation0 y %hich # really meant QappearedQ y some %holly
unkno%n process1( The (Pentateuchal term( 'ar%in is referrin! to here is the %ord (creation(1 The
context0 as ,rancis 'ar%in explains in his 2;;@ edition of his father(s letters0 %as that 'ar%in %as
%ritin! to thank )ooker for the loan of a revie% of a ook y Carpenter0 in %hich the anonymous
revie%er had spoken of (a creative force 1 1 1 %hich 'ar%in could only express in Pentateuchal terms
as the primordial form Qinto %hich life %as ori!inally reathedQ(1 *o%adays0 %e should dispense
even %ith the (ori!inally reathed(1 What is it that is supposed to have een reathed into %hatB
Presumaly the intended reference %as to some kind of reath of life0L
ut %hat mi!ht that meanB The harder %e look at the order et%een life and non:life0 the
more elusive does the distinction ecome1 -ife0 the animate0 %as supposed to have some sort of
virant0 throin! Euality0 some vital essence : made to sound yet more mysterious %hen dropped
into ,renchA elan ital4
-ife0 it seemed0 %as made of a special livin! sustance0 a %itch(s re%
called (protoplasm(1 Conan 'oyle(s Professor Challen!er0 a fictional character even more
preposterous than Sherlock )olmes0 discovered that the Earth %as livin!0 a kind of !iant sea urchin
%hose shell %as the crust that %e see0 and %hose core consisted of pure protoplasm1 Ri!ht up to the
middle of the t%entieth century0 life %as thou!ht to e Eualitatively eyond physics and chemistry1
*o lon!er1 The difference et%een life and non:life is a matter not of sustance ut of information1
-ivin! thin!s contain prodi!ious Euantities of information1 "ost of the information is di!itally
coded in '*&0 and there is also a sustantial Euantity coded in other %ays0 as %e shall see
#n the case of '*&0 %e understand pretty %ell ho% the information content uilds up over
!eolo!ical time1 'ar%in called it natural selection0 and %e can put it more preciselyA the non:
random survival of information that encodes emryolo!ical recipes for that survival1 Self:evidently
it is to e expected that recipes for their o%n survival %ill tend to survive1 What is special aout
'*& is that it survives not in its material self ut in the form of an indefinite series of copies1
Because there are occasional errors in the copyin!0 ne% variants may survive even etter than their
predecessors0 so the dataase of information encodin! recipes for survival %ill improve as time
!oes y1 Such improvements %ill e manifest in the form of etter odies and other contrivances
and devices for the preservation and propa!ation of the coded information1 On the !round0 the
preservation and propa!ation of '*& information %ill normally mean the survival and
reproduction of odies containin! it1 #t %as at the level of odies0 their survival and reproduction0
that 'ar%in himself %orked1 The coded information %ithin them %as implicit in his %orld:vie%0
ut not made explicit until the t%entieth century1
The !enetic dataase %ill ecome a storehouse of information aout the environments of the
past0 environments in %hich ancestors survived and passed on the !enes that helped them to do so1
To the extent that present and future environments resemle those of the past 7and mostly they do80
this (!enetic ook of the dead( %ill turn out to e a useful manual for survival in the present and
future1 The repository of that information %ill0 at any one moment0 reside in individual odies0 ut
in the lon!er term0 %here reproduction is sexual and '*& is shuffled from ody to ody0 the
dataase of survival instructions %ill e the !ene pool of a species1
Each individual(s !enome0 in any one !eneration0 %ill e a sample from the species
dataase1 'ifferent species %ill have different dataases ecause of their different ancestral %orlds1
The dataase in the !ene pool of camels %ill encode information aout deserts and ho% to survive
in them1 The '*& in mole !ene pools %ill contain instructions and hints for survival in dark0 moist
soil1 The '*& in predator !ene pools %ill increasin!ly contain information aout prey animals0
their evasive tricks and ho% to outsmart them1 The '*& in prey !ene pools %ill come to contain
information aout predators and ho% to dod!e and outrun them1 The '*& in all !ene pools
contains information aout parasites and ho% to resist their pernicious invasions1
#nformation on ho% to handle the present so as to survive into the future is necessarily
!leaned from the past1 *on:random survival of '*& in ancestral odies is the ovious %ay in
%hich information from the past is recorded for future use0 and this is the route y %hich the
primary dataase of '*& is uilt up1 But there are three further %ays in %hich information aout
the past is archived in such a %ay that it can e used to improve future chances of survival1 These
are the immune system0 the nervous system0 and culture1 &lon! %ith %in!s0 lun!s and all the other
apparatus for survival0 each of the three secondary information:!atherin! systems %as ultimately
prefi!ured y the primary oneA natural selection of '*&1 We could to!ether call them the four
The first memory is the '*& repository of ancestral survival techniEues0 %ritten on the
movin! scroll that is the !ene pool of the species1 .ust as the inherited dataase of '*& records the
recurrent details of ancestral environments and ho% to survive them0 the immune system0 the
(second memory(0 does the same thin! for diseases and other insults to the ody durin! the
individual(s o%n lifetime1 This dataase of past diseases and ho% to survive them is uniEue to each
individual and is %ritten in the repertoire of proteins that %e call antiodies : one population of
antiodies for each patho!en 7disease:causin! or!anism80 precisely tailored y past (experience(
%ith the proteins that characteriFe the patho!en1 -ike many children of my !eneration0 # had
measles and chickenpox1 "y ody (rememers( the (experience(0 the memories ein! emodied in
antiody proteins0 alon! %ith the rest of my personal dataase of previously vanEuished invaders1 #
have fortunately never had polio0 ut medical science has cleverly devised the techniEue of
vaccination for plantin! false memories of diseases never suffered1 # shall never contract polio0
ecause my ody (thinks( it has done so in the past0 and my immune system dataase is eEuipped
%ith the appropriate antiodies0 (fooled( into makin! them y the inHection of a harmless version of
the virus1 ,ascinatin!ly0 as the %ork of various *oel PriFe:%innin! medical scientists has sho%n0
the immune system(s dataase is itself uilt up y a Euasi:'ar%inian process of random variation
and non:random selection1 But in this case the non:random selection is selection not of odies for
their capacity to survive0 ut of proteins within the ody for their capacity to envelop or other%ise
neutraliFe invadin! proteins1
The third memory is the one %e ordinarily think of %hen %e use the %ordA the memory that
resides in the nervous system1 By mechanisms that %e don(t yet fully understand0 our rains retain a
store of past experiences to parallel the antiody (memory( of past diseases and the '*& (memory(
7for so %e can re!ard it8 of ancestral deaths and successes1 &t its simplest0 the third memory %orks
y a trial:and:error process that can e seen as yet another analo!y to natural selection1 When
searchin! for food0 an animal may (try( various actions1 Thou!h not strictly random0 this trial sta!e
is a reasonale analo!y to !enetic mutation1 The analo!y to natural selection is (reinforcement(0 the
system of re%ards 7positive reinforcement8 and punishments 7ne!ative reinforcement81 &n action
such as turnin! over dead leaves 7trial8 turns out to yield eetle larvae and %oodlice hidin! under
the leaves 7re%ard81 The nervous system has a rule that says0 (&ny trial action that is follo%ed y
re%ard should e repeated1 &ny trial action that is follo%ed y nothin!0 or0 %orse0 follo%ed y
punishment0 for example pain0 should not e repeated1(
But the rain(s memory !oes much further than this Euasi:'ar%inian process of non:random
survival of re%arded actions0 and elimination of punished actions0 in the animal(s repertoire1 The
rain(s memory 7no need for inverted commas here0 ecause it is the primary meanin! of the %ord8
is0 at least in the case of human rains0 oth vast and vivid1 #t contains detailed scenes0 represented
in an internal simulacrum of all five senses1 #t contains lists of faces0 places0 tunes0 social customs0
rules0 %ords1 6ou kno% it %ell from the inside0 so there is no need for me to spend my %ords
evokin! it0 except to note the remarkale fact that the lexicon of %ords at my disposal for %ritin!0
and the identical0 or at least heavily overlappin!0 dictionary at your disposal for readin!0 all reside
in the same vast neuronal dataase0 alon! %ith the syntactic apparatus for arran!in! them into
sentences and decipherin! them1
,urthermore0 the third memory0 the one in the rain0 has spa%ned a fourth1 The dataase in
my rain contains more than Hust a record of the happenin!s and sensations of my personal life :
althou!h that %as the limit %hen rains ori!inally evolved1 6our rain includes collective memories
inherited non:!enetically from past !enerations0 handed do%n y %ord of mouth0 or in ooks or0
no%adays0 on the internet1 The %orld in %hich you and # live is richer y far ecause of those %ho
%ent efore us and inscried their impacts on the dataase of human cultureA *e%ton and "arconi0
Shakespeare and Steineck0 Bach and the Beatles0 Stephenson and the Wri!ht rothers0 .enner and
Salk0 Curie and Einstein0 von *eumann and Berners:-ee1 &nd0 of course0 'ar%in1
&ll four memories are part of0 or manifestations of0 the vast super:structure of apparatus for
survival %hich %as ori!inally0 and primarily0 uilt up y the 'ar%inian process of non:random
'*& survival1
(#*TO & ,EW ,OR"S OR #*TO O*E(
'ar%in %as ri!ht to hed!e his ets0 ut today %e are pretty certain that all livin! creatures
on this planet are descended from a sin!le ancestor1 The evidence0 as %e sa% in Chapter 250 is that
the !enetic code is universal0 all ut identical across animals0 plants0 fun!i0 acteria0 archaea and
viruses1 The <>:%ord dictionary0 y %hich three:letter '*& %ords are translated into t%enty amino
acids and one punctuation mark0 %hich means (start readin! here( or (stop readin! here(0 is the same
<>:%ord dictionary %herever you look in the livin! kin!doms 7%ith one or t%o exceptions too
minor to undermine the !eneraliFation81 #f0 say0 some %eird0 anomalous microes called the
harumscaryotes %ere discovered0 %hich didn(t use '*& at all0 or didn(t use proteins0 or used
proteins ut strun! them to!ether from a different set of amino acids from the familiar t%enty0 or
%hich used '*& ut not a triplet code0 or a triplet code ut not the same <>:%ord dictionary : if
any of these conditions %ere met0 %e mi!ht su!!est that life had ori!inated t%iceA once for the
harumscaryotes and once for the rest of life1 ,or all 'ar%in kne% : indeed0 for all anyone kne%
efore the discovery of '*& : some existin! creatures mi!ht have had the properties # have here
attriuted to the harumscaryotes0 in %hich case his (into a fe% forms( %ould have een Hustified1
#s it possile that t%o independent ori!ins of life could oth have hit upon the same <>:%ord
codeB +ery unlikely1 ,or that to e plausile0 the existin! code %ould have to have stron!
advanta!es over alternative codes0 and there %ould have to e a !radual ramp of improvement
to%ards it0 a ramp for natural selection to clim up1 Both these conditions are improale1 ,rancis
Crick early su!!ested that the !enetic code is a (froFen accident(0 %hich0 once in place0 %as difficult
or impossile to chan!e1 The reasonin! is interestin!1 &ny mutation in the !enetic code itself 7as
opposed to mutations in the !enes that it encodes8 %ould have an instantly catastrophic effect0 not
Hust in one place ut throu!hout the %hole or!anism1 #f any %ord in the <>:%ord dictionary chan!ed
its meanin!0 so that it came to specify a different amino acid0 Hust aout every protein in the ody
%ould instantaneously chan!e0 proaly in many places alon! its len!th1 $nlike an ordinary
mutation0 %hich mi!ht0 say0 sli!htly len!then a le!0 shorten a %in! or darken an eye0 a chan!e in the
!enetic code %ould chan!e everythin! at once0 all over the ody0 and this %ould spell disaster1
+arious theorists have come up %ith in!enious su!!estions for special %ays in %hich the !enetic
code mi!ht evolveA %ays in %hich0 to Euote one of their papers0 the froFen accident mi!ht e
(tha%ed(1 #nterestin! as these are0 # think it is all ut certain that every livin! creature %hose !enetic
code has een looked at is descended from one common ancestor1 *o matter ho% elaorate and
different the hi!h:level pro!rams that underlie the various life forms0 all are0 at ottom0 %ritten in
the same machine lan!ua!e1
Of course %e cannot rule out the possiility that other machine lan!ua!es may have arisen
in yet other creatures that are no% extinct : the eEuivalent of my harumscaryotes1 &nd the physicist
Paul 'avies has made the reasonale point that %e haven(t actually looked very hard to see if there
are any harumscaryotes 7he doesn(t use the %ord0 of course8 that are not extinct ut still lurkin! in
some extreme redout of our planet1 )e admits that it is not very likely0 ut ar!ues : some%hat
alon! the lines of the man %ho searches for his keys under a street lamp rather than %here he lost
them : that it is a lot easier and cheaper to look thorou!hly on our planet than to travel to other
planets and look there1 "ean%hile0 # don(t mind recordin! my private expectation that Professor
'avies %on(t find anythin!0 and that all survivin! life forms on this planet use the same machine
code and are all descended from a sin!le ancestor1
(W)#-ST T)#S P-&*ET )&S GO*E C6C-#*G O* &CCOR'#*G TO T)E ,#CE' -&W O,
)umans %ere a%are of the cycles that !overn our lives lon! efore %e understood them1
The most ovious cycle is the dayGni!ht cycle1 OHects floatin! in space0 or oritin! other oHects
under the la% of !ravity0 have a natural tendency to spin on their o%n axis1 There are exceptions0
ut our planet is not one of them1 #ts period of rotation is no% t%enty:four hours 7it used to spin
faster8 and %e experience it0 of course0 as ni!ht follo%s day1
Because %e live on a relatively massive ody0 %e think of !ravity primarily as a force that
pulls everythin! to%ards the centre of that ody0 %hich %e experience as (do%n(1 But !ravity0 as
*e%ton %as the first to understand0 has a uiEuitous effect0 %hich is to keep odies throu!hout the
universe in semi:permanent orit around other odies1 We experience this as the yearly cycle of
seasons0 as our planet orits the sun1 L Because the axis on %hich our planet spins is tilted relative
to the axis of rotation around the sun0 %e experience lon!er days and shorter ni!hts durin! the half
of the year %hen the hemisphere on %hich %e happen to live is tilted sun%ards0 the period that
climaxes in summer1 &nd %e experience shorter days and lon!er ni!hts durin! the other half of the
year0 the period that0 at its extreme0 %e call %inter1 'urin! our hemisphere(s %inter0 the sun(s rays0
%hen they strike us at all0 do so at a shallo%er an!le1 The !lancin! an!le spreads a %inter suneam
more thinly over a %ider area than the same eam %ould cover in summer1 On the receivin! end of
fe%er photons per sEuare inch0 it feels colder1 ,e%er photons per !reen leaf means less
photosynthesis1 Shorter days and lon!er ni!hts have the same effect1 Winter and summer0 day and
ni!ht0 our lives are !overned y cycles0 Hust as 'ar%in said : and Genesis efore himA (While the